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HARVARD UNIVERSITY

Library of the

Museum of

Comparative Zoology

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PROCEEDINGS

OF THK

g0sl0n Somlg of |latural ^istorji*

VOL. XXIX.

WITH FORTY-ONE PLATES.

"^BOSTON:
PRINTED FOR THE SOCIETY.
1901.

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v^^

PrHLlSlIING COMMITTEE.

Charles 8. Minuj, Roland Thaxter,

Alpiiei's Hyatt, Jay B. Woodworth,

Glover M. Allen.

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CONTENTS OF VOL. XXIX.

Page
No. 1. — Proceedings of the Annual Meeting, May 3, 1899.

Report of the Curator, Alphbus Hyatt 1

Report of the Secretary and Librarian, Samtel Henshaw ... 14

Report of the Treasurer, £dwari> T. Boivfe 22

Officers for 1899-1900 24

List of members 26

By-laws 38

June, 1899.
No. 2. — Variation and Sexual Selection in Man. By Edwin Tennky

Brewster. July, 1899 46

No. 3. — Notes on the Reptiles and Amphibians of Intervale, New Hamp-
shire. By Glover M. Allen. July, 1899 63

No. 4. — Studies in Diptera Cyclorhapha. 1. The Pipunculidae of the

United States. By Garev i>e N. Hoigh. July, 1899. ... 77
No. 6. — Contributions from the Gray Herbarium of Harvard University.
New Series. — No. 17. By B. L. Robinson and J. M. Grbenman.

August, 1899 87

No. 6. — The Development of Penilia schmackeri Richai*d. By Mervin T.

Sri)LER. (3 Plates.) October, 1899 109

No. 7. — List of Marine Mollusca of Coldspring Harbor, Long Island, with

descriptions of one new Genus and two new Species of Nudibranchs.

By Francis Noves Balch. (1 Plate.) October, 1899. ... 133

No. 8. — The B1(mk1 Vessels of the Heart in Carcharias, Raja, and Amia.

Bv G. H. Parker and Frei>erica K. Davis. (3 Plates.) October,

1899. 103

No. 9. — The Oceurrt»nce of Fossils in the Roxburj' Conglomerate. By

Henry T. Birr and R^uieht E. Birke. (1 Plate.) April, 1900. 179
No. 10. — On a hitherto unrecognized fonn of bUxMl circulation without
capillaries in the organs of the Veitebrata. By Charles SEiMiwuK

Minot, LL. D. April, HKH) 186

No. 11. — A Revision of the Systematic Names employed by Writers on
the Morpholo^' of the Acmaeidae. By M. A. Wii.lcox, Ph. D.

April, liHK) 217

No. 12. — Proceedings of the Annual Meeting, May 2, 1900.

Re|X)rt of the Curator, A LPiiEi-s Hyatt 223

Report of the Secretary- and Librarian, Charles F. Batcheldbr 234

Report of the Treasurer, Ei»ward T. Borv6 239

May, 1900.
No. 13. — The Embryonic History of Imaginal Discs in Melophagus ovinus
L., together witli an Account of the Earlier Stages in the Develop-
ment of the Insect, By H. S. Pratt, Ph. D. (7 Plates.) June,

19(>f) 241

No. 14. — Glacial Erosion in France, Switzerland and Norway. By

William Morris Davis. (3 Plates.) July, 1900. . . .273

No. 16. — Echinoderms from Puget Soiuid : Ol)servations made on the
Echinoderins collected by the parties from Colimibia University, in
Puget Sound in 1890 and 1897. By Hubert Lyman Clark. (4

Plates.) May, 1901 323

No. 16. — Bermudan Echinoderms. A Report on Observations and Collec-
tions made in 18m). By Hubert Lyman Clark. May, 1901. . 339
No. 17. — Proceedings of the Annual Meeting, May 1, 1901.

Report of the Curator, Alpheis Hyatt 347

Report of the Secretary and Librarian, Charles F. Batchelder . 358

Report of the Treasurer, £dwari> T. Bouvfe 304

Officers for 1901-1902 300

List of Members 307

June, 1901.
No. 18. — The Polychaeta of the Puget Souna Region. By Herbert

Farlin Johnson. (19 Plates.) August, 1901 381

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Vol. t% No. I,

l'lt*iCKKnrNr}H OK TIIK ANNTAL MKKTINO. MAY :i, IHOO

BOSTON
PHINTEI> FflR TITE SOOIETT
June, 1899-

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No. 1. — Proceedings of the Annual Meeting^ May 3, 1899.
Report of the Curator, Alpiieus Hyatt.

The death of Mr. John Cummings of Woburn removes a member
to whom this Society is indebted for valuable time freely given for
many years in its Council and for much actual work done in its
Museum, especially in the botanical department. While his pecun-
iary means were ample, he was as generous in giving aid with
them as with his brain and hands. For a long time he carried on
the botanical department, working in it himself and paying the
salary of an assistant; he also maintained another assistant in the
3Iuseum, and at the same time supported the Teachers' school of
science. His services in the botanical department were described in
detail in my report for 1898 on the occasion of his retirement from
that department which he had sustained from 1873 to 1898 and
which he had succeeded in placing in excellent. condition. Another
monument to Mr. Cummings's generosity is the collection of European
fossils filling Room H. This is the Eser collection, which is entirely
his gift, and is one of the most famous of the older and smaller
European collections. This collection is a great prize for any
museum since it possesses very rare and valuable specimens and is
especially suitable for the purposes of our educational series. The
Teachers' school of science owes its foundation to Mr. Cummings
and arose in the Council in conseqlience of his offer to support a
series of lectures for teachers. The Curator immediately accepted
this offer, and the school began in the following autumn. These
are only his principal claims to our remembrance and gratitude,
since it is impossible to take notice of the thousand and one ser-
vices to the cause of science which he gave so generously and with-
out expectation of credit or reward.

In his last annual report the Curator called attention to the
necessity of doing something to perpetuate the office of Guide to
the Museum. This has been held by Mr. Grabau and has become
really a free lectureship, that has not only made our collections
more instructive to the public but also interested a number of
persons in the study of natural history and led to the giving of

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2 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

many lessons on the sea- shore and in our laboratory or lecture room.
This has greatly increased our usefulness to the community, and
some means should be found to continue it.

An appeal was also made for aid in the botanical department.
This is still cared for by Miss Carter but largely as a voluntar}*
service, since only somewhat more than is sufficient to meet her
expenses is now paid her.

Another part of the same report set forth the claims of the New
England collection in our Museum, and the great need that exists
of separating this from the educational and general systematic
collections. The New England collections are scattered in the
different departments and should be brought together in order to
give a complete exposition of the natural products of our own
neighborhood. The need of exhibiting in connected succession
the mineralogy, geology, botany, and zoology of New England
was fully demonstrated during the past summer when the American
association for the advancement of science met in our building
and in that of the Institute of technology and many of its members
visited our Museum. If we had been able to throw open to them
a series of collections showing such a connected history of the
natural products of New England and of the geology of this region,
it would have been a revelation of the teaching j)ower of collections
and might have had a far-reaching influence upon other museums in
this country.

The Curator has always held that a New England museum
should be brought together in this way within our building, and
the whole be placed in order before any General guide could be
effectively written. This year, however, the need for some guide
to explain the apparent confusion to scientific visitors became urgent,
and he wrote a small pamphlet of forty-seven pages, accompanied
by a diagram of the building. This appeared as a second edition
of a General guide written and published on the celebration of the
fiftieth anniversary of this Society, but it is in reality new.

Teaching ix the Museum.

The operations of this department have ceased for want of funds,
as has been noted above.

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HYATT: REPORT OF THE CURATOR.

Dynamical Zoology.

Considerable work on this collection has been done. The differ-
ent series have been rearranged and general descriptive labels
prepared by the Curator assisted by Miss Bryant. The large relief
map of Oahu shows with the aid of colored pins the migrations of
the species of three of the principal genera of Achatinellidae. The
genera are represented by pins with heads of different colors, and
the species are represented by different numbers and labels on the
pins. In this way the migrations of the species may be followed
along the mountain ridges of the island, and the correlation of
these movements with topography is brought out more clearly than
by the use of the shells themselves. A valuable and interesting
series of lamprey eels has been given to this collection by Prof.
S. H. Gage of Cornell.

Mineralogy and Geology.

During the summer Professor Crosby gave considerable time to
the geological department of the Museum, putting the specimens
on exhibition in more perfect order, labeling new specimens, etc.,
and he also labeled and arranged, as well as the limited space now
available would permit, the illustrative specimens and maps pre-
pared for Parts 1 and 2 of the Boston Basin work, and for Part 3
so far as that was then completed. The same assistant has also
completed a much needed revision of the general collection of min-
erals and incorporated with the mounted s})ecimens on exhibition
all of the materials that have been accumulating in the past ten
years. Professor Crosby has also personally paid for the necessary
clerical assistance employed in this work.

During the meeting of the American association for the advance-
ment of science he presented before Section E an outline of the
history of the Blue Hills complex, and abstracts of this paper
appeared in the American geologist, in Science, and in the Pro-
ceedings of the American association. He also conducted seven
parties of geologists to points of local interest, including the Blue
Hills, and had the satisfaction of learning that his views concerning
the geology of this exceedingly difficult region were endorsed by
competent authorities.

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4 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

Another gratifying result of going over the ground with these
geologists was the offer of further cooperation. Professor Crosby's
work is necessarily mainly areal and structural, and although he
has himself paid for a number of chemical analyses and has received
additional assistance from several of his students, it was very de-
sirable that this should be supplemented, especially for the igneous
rocks, by microscopic and chemical work. Dr. White of Columbia
has very kindly done considerable microscopical work on the Blue
Hills rocks, and Dr. A. S. Eakle of Harvard has kindly supple-
mented this by investigations of the plutonic rocks, especially on
chemical lines. The intricate nature of the problem of the geology
of this region has been the cause of the successive delays in com-
pleting the work, and it would still have remained unfinished if
Professor Crosby had not this year given an extraordinary amount
of time to this purpose.

The manuscript of Part 3, *' The Blue Hills complex," is now in
the hands of the Secretary, 375 pages, with 8 plates and 26 figures.
Two chapters which Mr. Grabau is writing, one on the fossils and
one on Lake Bouv6, an extinct glacial lake, noticed in the report
for 1895-96, are in the hands of the author, but are stated to be
practically finished. An important accession of new fossils lately
received, including the Sears collections from Nahant and Mr.
"W. W. Dodge's collections from Braintree, has prevented the com-
pletion of the descriptions of the fossils in time for this report, but
it is gratifying to notice that these accessions have about tripled
the materials for investigation.

The work on the Neponset Valley, Part 4, of the Boston Basin
work, has been actively pressed. Outside the areal and structural
work done by Professor Crosby, Dr. Florence Bascom, of Bryn
Mawr college, accepted an invitation to contribute by studying the
volcanic rocks, and Mr. F. C. Ohm of the U. S. geological survey
has also kindly taken the work on the thin sections, while Professor
Crosby has arranged for a sufiicient number of chemical anal3\ses
to supplement Miss Bascom 's investigations.

Professor Crosby ends his report to the Curator as follows : '* The
willingness of other geologists to cooperate with me is a great
encouragement and leads me to hope that the ideal end of this work
may be realized ; viz : an illustrate<l handbook of the local geology of
our own neighborhood accompanied by a complete collection on
exhibition in suitable rooms. This should not only be creditable to

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HYATT: REPORT OF THE CURATOR. 5

the Society and of great educational value, but also stand the test
of close and critical comparison with facts in the field,"

That this hope is not too highly aimed is shown by the fact, that
it is even now in large part realized. This can be demonstrated by
the accumulated collections and maps, by the willingness of able
observers to cooperate in the laboratory and in the field, and also
by a published opinion emanating from the highest authority. The
Hon. C. D. Walcott, Director of the U. S. geological survey and
the leading investigator of Cambrian rocks, wrote in 1871 of Pro-
fessor Crosby's work in his Correlation of the Cambrian, Bull. U. S.
geol. surv., no. 81, p. 268, as follows: " The description of the Cam-
brian rocks of the Boston Basin by Professor W. O. Crosby is the
most recent and the most thorough of any we have, and the sum-
mary of our present knowledge of them is taken mainly from his
paper."

Synoptic Zoology.

Mrs. J. M. Arms Sheldon has contributed by the purchase of
suitable specimens for this department, having drawings framed,
etc., at her own expense, as well as by the donation of far more
personal work than the time called for by the duties of her position
as assistant in the Museum.

She spent considerable time in the spiing and summer in making
the whole collection more presentable and in placing on exhibition
most of the fine specimens and drawings that have been slowly accu-
mulating during several years past, so that the larger part of the
Invertebrata now present a far better aspect than at the beginning
of the present official year. About 314 specimens and 61 plates
containing 266 figures were added to the collection, 172 of which
were mounted by Mrs. Sheldon.

The worms have been completed, 67 drawings having been
selected and arranged in plates, 12 tablets of mounted specimens
added to this section of the collection, and about 95 pages of manu-
script written. Considerable work has also been done upon the
Brachiopoda and Pelecypoda, and upon the arrangement of Tuni-
cata. Eighty specimens mostly Brachiopoda have been mounted
and placed on exhibition ; also eight genera of Tunicata and two
of Cephalopoda. Considerable additions have also been made to
the text of the Guide in the Actinozoa and Brachiopoda and some-

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(> PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

thing on Pelecypoda. A considerable number of fossil Gasteropoda
have also been selected from the general collections, mounted, and
placed on exhibition. Mrs. Sheldon has also described about fifty-
eight species of Crustacea and selected and mounted a limited num-
ber of specimens.

Miss Martin has been as usual more or less occupied in making
colored drawings for this collection, especially during the summer.
The labeling of the tablets has been done by Miss Bryant and also
the preparation of some of the fossils.

Botany.

Less work than in previous years has been done u})on the herba-
rium, owing in part to the unfortunate sickness of the assistant.
Miss Carter. She nevertheless continued through a large part of
the past official year the work u])on the labeling of the Lowell col-
lection and systematic arrangement of the duplicates and the j)oi-
soning of the plants. During the summer she also looked over and
cleaned the specimens on exhibition and placed in position a series
of shelf labels indicating the different divisions of plants, and Miss
Bryant a.ssisted her by making these labels. Thirty species of North
American lichens were added by purchase from Cummings, Seymour,
and Earle. Nineteen persons have been permitted to consult and
study in the herbarium.

Palaeontology.

Miss Bryant has this year worked upon the renovation of the
tablets and specimens in the European room, especially in Oolite,
and has rewritten about four hundred labels. She has also worked
over all of the American Jurassic and Cretaceous fossils and has
cleaned from the matrix and mounted a number of Silurian fossils
of the American collection from Anticosti, and a collection of fos-
sils from the St. John's group has also been cleaned and mounted
by the same assistant. A number of Devonian corals purchased
from Mr. Greene have been identified and labeled, and the Vertebrata
from the Cretaceous freshly labeled. The same assistant revised
the work previously done by other assistants upon the Trenton and
Niagara fossils and relabeled them. She also finished the Carboni-
ferous fossils including the Brachiopoda, Lamellibranchiata, Gastero-
poda, and Cephalopoda.

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HYATT : REPORT OF THE CURATOR.

MOLLUSCA.

The large collection purchased of Rev. J. T. Gulick last year and
described in the report of 189H has been completely catalogued and
every lot labeled, each shell having been also numbered. The col-
lections of Achatinellidae previously mentioned have been still
further increased by the loan of eight thousand shells belonging to
Mr. C. M. Cooke, Jr., of Honolulu, making in all about thirty
thousand shells of this family at present available for study. The
Curator has found it essential to trace as far as practicable the
migrations of the Achatinellae from island to island, and has made
good progress during the past year, having reviewed most of the
genera that are found in islands outside of Oahu. A small but
valuable collection of Achatinellidae from the island of Molokai
that filled a number of blanks in our own collection has been given
us by Dr. W. P. Wesselhoeft. Miss Martin completed the work of
labeling the duplicate Gasteropoda having no locality labels, and
these were in part given away to the public schools and in part
exchanged for invertebrates collected in the West Indies by Mr.
C. J. Maynard. Miss Martin has made a complete series of colored
drawings of the animals of Pteropoda and arranged these in com-
pany with a series of the shells purchased from Mr. Sowerby.

The death of Mr. Edward W. Roper, a member who had taken a
great interest in this collection and was practically an assistant in
this department, was a severe loss. He had, in the short time that
he was connected with us before his health obliged him to seek
refuge in southern California, done much efficient work, and had
also planned the revision of all our land shells. His entire con-
chological collection was bequeathed to us and is now in our pos-
session, with the exception of the Cyrenidae. These are in the
hands of Mrs. Roper, who will deliver them to us as soon as prac-
ticable after her return to the east. A detailed re])ort on this
collection will be made after these have been received and the
whole collection has been catalogued. This work has been begun
by Miss Martin.

Ckustacea.

The work of restoring the faded labels of alcoholic preparations,
begun by Miss Martin last year, was renewed this year, and will be

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8 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

continued during the coining year. Professor Kingsley of Tufts
college, to whom our collection of Amphipoda was loaned for
investigation, reports that about one fourth of the specimens have
been named and that the whole collection is in good condition.

Birds and Mammals.

The New England collection of birds was removed from the cases
during the early part of this official year and the new backs put
into the cases with its new shelving brings the specimens close to
the glass where they can be readily seen and examined. The
specimens were removed and returned to their places by Miss
Martin, and the arrangement was subsequently revised by Mr.
Batchelder.

The Mammalia were also removed and the cases in the main
gallery immediately adjoining the entrance to Room N were
deepened. This enabled Miss Martin to place the collection of
New England mammals outside of Room N and thus make suffi-
cient space for the extension of the collection of birds within that
room. The arrangement of the mammals also was revised by Mr.
Batchelder.

A railing case of improved pattern was built early in the summer
along the east side of the upper gallery for the accommodation of
the larger specimens of birds' nests and eggs promised by the
Nuttall ornithological club. Mr. Batchelder removed from their
cases all of the old collection of birds' nests and eggs, which had
for some years been in bad condition, and cleaned and rearranged
them, weeding out undesirable material during the process. A part
of the donation of the Nuttall club has been received and placed
in the cases by Mr. Batchelder. This makes a fine appearance,
and the sincere thanks of the 'Society are due to the members of
the Club for a donation which has so greatly improved this part
of our New England collections, and to Mr. Batchelder for the
expense and trouble incurred in appropriately mounting these
handsome specimens. Unfortunately Mr. 15atchelder met with a
serious accident a few weeks since, and this has stopped for the
present the active work he was doing in the collections under his
charge, but, as this report shows, his efforts this year had already
laid the Society under obligations which should be gratefully
acknowledged.

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HYATT : REPORT OF THE CURATOR. 9

During the summer the Curator succeeded in obtaining a rare
species of whale, Meaoplodon hidens^ noticed at the meeting of
November 2, 1898. The bones of this specimen are now being
prepared for the collection.

Laboratory.

The room in our basement has been this year used as in previous
years by the classes of the Boston univei-sity and Teachers* school
of science. A number of new diagrams have been made for use
in the laboratory and the specimens have been looked after by the
Curator and Miss Martin.

Remarks.

A much larger amount of work upon the collections not reported
upon above has been done this year than usual, especial efforts
having been made to bring into order and work up our miscella-
neous alcoholic and dry materials. It should also be noted here
that these annual reports take no notice whatever of this sort of
work nor of similar work upon the various collections, such as the
general inspection and repair of dried specimens, the poisoning of
the same twice in each year, the inspection of alcoholic sj)ecimen8
and the refilling of bottles once in each year, the reception and
preparation of specimens, and other items of daily routine.

The Museum has been visited during private days by 340 pupils
and teachers, representing 12 schools, all of whom have as heretofore
been admitted without charge. It should also be noted here that
this year as in previous years several artists have been allowed to
draught specimens or study them in the Museum on closed days
free of charge.

Teachers' School of Science.

Mr. Grabau has continued his excursions and lectures upon zool-
ogy. The Satm-day field courses have been regularly carried on.
Four excursions were made in May and June, 1898, in addition to
those reported upon in the last annual report. These took in the
more important localities on our coast and the fresh-water ponds of
West Cambridge. A three days' excursion to Wood's IIoU was

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10 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

also made and notwithstanding the length of time consumed and
the expense of such a prolonged trip, this was taken by ten persons.
The days were spent in the general study of the fauna of the shores
and the evenings in surface work. Reports and theses on the speci-
mens found and preserved were subsequently prepared, each student
selecting a distinct group of animals. Another longer excursion of
ten days was made to Oouldsbo rough on Frenchman's Bay, Maine.
Several boats together with dredging apparatus were placed at the
command of this party. The habits of marine animals were the
principal subject for observation on this trip. As on previous trips
an effort was made to collect extensively, and lists of the species are
in preparation as part of the theses of the members of the clans.
Several members of this class made an indei)endent excursion of
several weeks' duration to the island of Monhegan and collected a
large number of marine invertebrates, some of which have not been
heretofore reported from this part of the coast. The autumn course
of field lessons in zoology also given by Mr. Grabau began Septem-
ber 10, and ten lessons were given ; but owing to the exceptional
inclemency of the weather, eight of these were necessarily trans-
formed into laboratory exercises. The average attendance was
twenty.

The Hale house natural history club, founded by members of
these classes, still continues to work in cooperation. All remunera-
tion ceased with the close of the spring courses, and since then
Mr. Grabau has been conducting the instruction without pay,
excepting in so far as the members of the classes have defrayed his
traveling expenses and the cost of circulars and correspondence.
Unless some substantial support can be obtained for this work, it
must cease, and the fruit of years of preparation as well as several
years of direct work will be lost. A few hundred dollars per year
would enable us to carry on these classes which must eventually
exercise a large influence upon our efforts to investigate and excite
a general interest in the natural history of New England.

Our work upon the geology of this region is in full swing with
two able men, one constantly investigating and publishing, the
other at the head of a large, enthusiastic, and increasing class of
teachers in the public and private schools.

The zoology has also been successful and will do quite as much
for the biological side of our work, if it meet with equal patronage.

The spring work in zoology has been begun with one regular

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m'ATT: REPORT OF THE CURATOR. 11

excursion to Nahant and a number of informal trips, and will be
continued for the present.

Professor Barton had arranged to give ten field lessons in geology
to the pupils of the Boston normal school but the exceptionally
bad weather in the autumn of 1898 reduced these to four. This
class consisted of twenty young ladies and was conducted without
remuneration. The indifference of the authorities to the continu-
ance of this course, in spite of the exertions of the teacher in charge
and of the head of the school to obtain an appropriation for this
purpose, exhibits quite j)lainly the estimate in which science is held
by the government of the public schools. Professor Barton states
that these pupils, all of whom are to be teachers in the })ublic
schools, come to him exceedingly well prepared for the work by
their previous training and are very enthusiastic and successful, but
as he has now been carrying these courses for several years without
remuneration he will be unable to continue after the series of
lessons now being given is finished. The spring course has begun
and will be reported upon next year.

The field lessons in geology in the autumn of 1898, altliougli no
longer supported by the Lowell fund, were conducted voluntarily
by Professor Barton in order that the teachers who had counted
upon having them should not be disappointed and in the hope that
means would be found to keep this important part of the work of
the school from being given up altogether. Luckily our appeals
for help were in this case answered by a generous friend of the
Society, who has promised a sum sufficient to cany on these courses
through the spring and autumn of 1899.

The autumn course began September 17, 1898, and ended
November 19, ten lessons having been given. The whole number
of applications was 110 and the average attendance notwithstand-
ing the b«id weather was twenty-seven. There was but one fair day
out of the ten Saturdays upon which lessons were held. Most of
these lessons occupy one half day but some, like those to Marble-
head, Rockport, Haverhill, Fitchburg, require a whole day and
those to Iloosac Tunnel and Mt. Ilolyoke took three days each.
The spring course of 1899 has begun and will be noticed fully in
the next annual report.

A party of seventeen under Professor Barton's direction visited
Nova Scotia during the summer of 1898, spending about three
weeks in making a study of the geology, mineralog}% mining, and

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12 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

natural scenery of that country. Among the prominent points
visited were the coal mines at the Joggins, the iron mines at
Londonderry, the gold mines at Montagu, the gypsum quames at
Windsor, the famous mineral localities at Partridge Island, Wason's
Bluff, and Cape Blomidon, and the beautiful scenery around Hali-
fax and the Annapolis Valley. Many courtesies were shown the
party by the managers of the different mines and by others. The
members of the class were charged with the teacher's expenses
and a fee of five dollars each. A similar excursion into western
Massachusetts and westwards as far as Niagara Falls has been
planned for the coming summer.

Lowell Fbee Courses.

The regular spiing course of ten field lessons in geology car-
ried on by Professor Barton began April 23, 1898, and ended
June 25. Bad weather interfered with the work and reduced the
attendance to less than it ever has been before. While in 1897 for
the same course it was 34, this spring the average was only 10.

The Curator regrets to announce that the Trustee of the Lowell
institute having decided to discontinue the out-of-door work of the
Lowell free course in the Teachers' school of science, will contrib-
ute in future only to the support of lectures during the winter. To
this determination he has been led by considerations of general
policy which in no way reflect upon the value or success of the
field courses.

It would be difficult to overstate the obligations of this Society
to Mr. Augustus Lowell. This gentleman as Trustee of the Lowell
institute has carried on for many years past most of the les-
sons given by the Teachers' school of science and preceding this
time gave freely towards the maintenance of courses of free public
evening lectures. He assumed the support of the field lessons in
geology in 1890, and they have since then been a part of the work
of the Lowell institute and have thus been able to build up results
that ought to secure for them some permanent foundation.

A new four years' course (1 20 hours) in geology under Profes-
sor Barton began during the past winter. This course like that
which preceded it will embrace at least fifteen lessons of two hours
each, thirty hours for each year of the course. The subjects are to

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HYATT: REPORT OF THE CURATOR. 13

be as follows : mineralogy, lithology and dynamic geology, struc-
tural geology, and historical geology.

The first year's course on mineralogy began on December 5,
1898, and ended on April 15, 1899. This included one extra
lecture which was given by special request at the end of the regular
lessons and the final examination, making seventeen exercises, in
all thirty-four hours of instruction instead of the required thirty
hours. The entire number of applications received for this course
was 145, and special provision was made to accommodate 130 of
these. The unusual prevalence of sickness during the winter
caused the loss of several from the class and a few also withdrew as
they found the work more difticult than they could attend to in
connection with the necessary work in their schools. The average
attendance for the course was 106. Of these 72 took all the exam-
inations, including the final. The instruction included one intro-
ductory lecture upon chemistry, four upon crystallography, and ten
upon mineralogy proper. They were given by means of lectures,
supplemented by a complete series of minerals illustrating the com-
moner species, about 130 in number. Each two members of the
class had one tray between them, containing all the species dis-
cussed at any single lesson, and this enabled each member of the
class to study with the specimens directly in hand. A complete
set of notes was also supplied each member of the class, for which a
cost price was charged. At each lesson except the first and the
last the first half hour was devoted to an examination covering all
the ground previously passed over, and at the end of the course a
final examination of three hours was given which was so arranged
as to present a concise r4sum6 of the whole subject.

Dr. R. W. Greenleaf gave a course of fifteen lessons of two hours
each upon the principles of the classification of flowering plants.
One hundred species of native plants were collected, dried, mounted,
and divided into sets by Miss E. B. Bryant, and each student was
provided with one of these. Fresh material was also purchased
when required for the use of the class. The class was especially
mdebted to Miss Helen Sharp, formerly an assistant in this course,
for the use of her collection of water-color drawings of 901 sheets,
774 of which represent American plants, the actual number of
American species being 649. These drawings will shortly be
assembled and exhibited in our laboratory, where they can be seen
as a whole and properly inspected. This free public exhibition will

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14 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

take place between the 9th and 13th of May, and will be open daily
from 12 to 4.30. There were forty persons in the class and the
average attendance was twenty-seven. Twenty-one persons took
the examination, out of which number nine passed with honor,
seven with credit, four simply passed, and one failed.

The Curator gave the fourth year of a five years' coui-se in
zoology, consisting this year of sixteen lessons of two hours each.
The subjects were Myriopoda, Arachnozoa, and Insecta through
Coleoptera, leaving the Lepidoptera and other so-called higher
orders to be treated next year. The number of tickets issued was
forty-five and the average attendance twenty-five. The excessively
bad weather had a depressing effect upon this course as upon most
others. Thirteen only took the final examination and all of these
passed.

REPORT OF THE SECRETARY AND LIBRARIAN, SAMUEL
HENSHAW.

Membekship.

During the past year eighty-four Corporate members have been
elected by the Council. Nine Corporate members have become
Life members. One Honorary member, James Hall, has died.
Three Corresponding members, George Baur, Oliver Marcy, and
O. C. Marsh, have died.

From the list of Corporate members we have lost by death five
names, John Cummings, Edraond E. Fernald, James I. Peck,
Henry P. Quincy, and Edward W. Roper.

Two Corporate members have resigned, and the names of four-
teen have been stricken from the list for non-])ayment of dues.

The membership of the Society, corrected to May 3, 1899, con-
sists of 10 Honorary, 139 Corresponding, and 420 Corporate mem-
bers, a total of 569. There are 17 Patrons.

The numl>er of Corporate members reported last year was 363 ;
twenty-three is the greatest numl>er elected in any of the antecedent
seven years, and for the essential increase this year the Society is
indebted mainly to the abundant faith and efticient work of its
President, Professor Minot.

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HENSHAW: REPORT OF SECRETARY AND LIBRARIAN. 15

The Dames of the Corporate members elected and the dates of
their election are as follows : —

Arthur Amory, Feb. 15, 1899.

S. Reed Anthony, Feb. 15, 1899.

Francis Blake, Feb. 15, 1899.

Mrs. T. M. Brewer, Feb. 15, 1899.

Shepherd Brooks, Feb. 15, 1899.

Edward I. Browne, Feb. 15, 1899.

Arthur T. Cabot, Mar. 29, 1S99.

William B. Cabot, Apr. 26, 1899.

Alvin Carl, Mar. 29, 1899.

Mrs. J. B. Case, Mar. 29, 1899.

John S. Clark, Mar. 29, 1899.

Collier Cobb, Feb. 15, 1899.

Miss Helen Collamore, Feb. 15, 1899.

T. Jefferson Coolidge, Jr., Feb. 15, 1899.

Charles U. Cotting, Feb. 15, 1899.

Miss S. H. Crocker, Feb. 15, 1899.

Miss Ada Dana, Oct. 19, 189><.

Andrew McF. Davis, P\»b. 15, 1899.

James C. Davis, Mar. 29, 1899.

Gordon Dexter, Feb. 15, 1899.

Thomas Doliber, Mar. 29, 1899.

Richard S. Dow, Feb. 15, 1899.

Raymond B. Earle, Feb. 15, 1899.

G. B. Eisenhard, Oct. 19, 1898.

Mrs. J. W. Elliot, Mar. 29, 1899.

George W. Fitz, Feb. 15, 1899.

Charles F. Folsom, Feb. 15, 1899.

Eugene N. Foss, Mar. 29, 1899.

Charles Fry, Mar. 29, 1899.

George M. Garland, Feb. 15, 1899.

George W. Gay, Feb. 15, 1899.

Harold B. Goodrich, Mar. 29, 1899.

Mrs. Mary T. Gorham, Feb. 15, 1899.

Francis C. Gray, Feb. 15, 1899.

Mrs. Caroline W. Greenough, Mar. 29, 1899.

Charles P. Greenough, Feb. 15, 1899.

Elisha H. Gregory, Jr., Oct. 19, 1898.

Miss Minna B. Hall, Mar. 29, 1899.

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16 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

Francis R. Hart, Feb. 15, 1899.
Charles E. Hellier, Feb. 15, 1899.
Mrs. Augustus Hemenway, Feb. 15, 1899.
Joseph P. B. Henshaw, Feb. 15, 1899.
Hibbert \V. Hill, Mar. 29, 1899.
Robert C. Hooper, Feb. 15, 1899.
Theodore Hough, Mar. 29, 1899.
John E. Hudson, Feb. 15, 1899.
Willard P. Hunnewell, Mar. 29, 1899.
Charles E. Inches, Feb. 15. 1899.
Charles F. Jenney, Oct. 19, 1898.
Miss Marian H. Judd, Feb. 15, 1899.
William Lawrence, Feb. 15, 1899.
George W. Lee, Dec. 21, 1898.
William C. Loring, Feb. 15, 1899.
Charles Lowell, Apr. 26, 1899.
Charles P. Lyman, Apr. 26, 1899.
Vernon F. Marsters, Mar. 29, 1899.
Asa E. Mattice, Mar. 29, 1899.
Laurence Minot, Feb. 15, 1899.
William Minot, Feb. 15, 1899.
William J. Moenkhaus, Dec. 21, 1898.
Miss Margaret W. Morley, Feb. 15, 1899.
Elisha W. Morse, Dec. 21, 1898.
Mrs. Edith J. Nichols, Mar. 29, 1899.
Edgar W. OUve, Oct. 19, 1898.
Edward C. Perkins, Feb. 15, 1899.
John C. Phillips, Oct. 19, 1898.
Dudley L. Pickman, Feb. 15, 1899.
David Pingree, Feb. 15, 1899.
William H. Ruddick, Dec. 21, 1898.
Mrs. T. E. Ruggles, Oct. 19, 1898.
Dudley A. Sargent, Mar. 29, 1899.
Alfred L. T. Schaper, Feb. 15, 1899.
Frederic F. Smith, Feb. 15, 1899.
John E. Thayer, Feb. 15, 1899.
Augustus L. Thorndike, Mar. 29, 1899.
Miss Mary M. Webster, Feb. 15, 1899.
Clarence M. Weed, Oct. 19, 1898.
William P. Wesselhoeft, Feb. 15, 1899.

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HEXSHAW; REPORT OF SECRETARY AND LIBRARIAN. 17

Henry M. Whitney, Feb. 15, 1899.
George Wigglesworth, Feb. 15, 1899.
Guy M. Winslow, Feb. 15, 1899.
Edward S. Wood, Mar. 29, 1899.
Miss Elvira Wood, Dec. 21, 1898.
Frederick A. Woods, Dec. 21, 1898.

Meetings.

Fourteen regular meetings including the Annual meeting have
been held during the year; one Special meeting has been held.
The attendance shows an average of 55 plua^ the largest at any one
meeting being 144, the smallest 24 ; the largest last year was 352,
the smallest 24.

Twenty-six communications have been made by twenty-two
persons ; of the twenty-two persons seven have not previously
spoken at our meetings.

Twelve papers have been presented by title.
The substitution of electricity for gas in the lecture room and
the availableness of the lantern for even a modicum of illustration
are features that should add to the success of future meetings.
The Society and especially the Secretary are indebted to Dr.
R. T. Jackson for his kindness in serving as recorder at meetings
in January and February.

The meetings, attendance, and communications have been as
follows : —
May 4, 1898. Annual meeting. Forty-two persons present.

Reports 'of the Curator, Secretary, Librarian, Treasurer, and

Trustees.
Mr. J. Edmund Woodman. Geological history of the gold-
bearing slates of Nova Scotia.
Mr. M. L. Fernald. The genus Antennaria in New England.

(By title.)
Prof. C. S. Minot. On the veins of the Wolffian bodies in the

pig. (By title.)
Dr. P. P. Calvert. The odonate genus Macrothemis and its

allies. (By title.)
Mr. T. D. A. Cockerell. The North American bees of the
genus Prosapis. (By title.)

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18 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

May 18, 1898. General meeting. Thirty-two persons present.
Dr. T. A. Jaggar, Jr. The Absaroka range of the Rocky
Mountains.
November 2, 1898. General meeting. Thirty-seven persons present.
Prof. Alpheus Hyatt. A rare whale from the Massachusetts

coast.
Dr. R. T. Jackson. Localized stages in growth in plants and

animals.
Dr. G. N. Calkins. Some hydroids from Puget Sound. (By

title.)
Mr. Reginald Heber Howe, Jr. North American wood frogs.
(By title.)
November 16, 1898. General meeting. Eighty-five persons present.
Prof. W. Z. Ripley. Racial characteristics of the Jews — a
study in physical anthropology.
December 7, 1898. General meeting. Forty-five persons present.
Dr. G. H. Parker. On the coronary circulation in some fishes.
Prof. M. A. Willcox. On the occipital region of the head in

the European trout.
Prof. J. S. Kingsley. Ear-bones of mammals.
Prof. C. S. Minot. Prof. O. van der Stricht's demonstration

of the human ovum.
Prof. C. S. Minot. Some classical embryological monographs.
December 21, 1898. General meeting. Forty-three persons pres-
ent.
Prof. Alpheus Hyatt. Evolution and migrations of the land
shells of the Hawaiian Islands.
January 4, 1899. General meeting. Thirty-six persons present.
Mr. J. B. Wood worth. The geology and geography of the
Richmond area in Virginia.
January 18, 1899. General meeting. Forty- four persons present.
Dr. Frank Russell. An account of the Apache festival of San
Antonio.
February 1, 1899. General meeting. Seventy-six persons present.
Dr. R. A. Daly. A geological tour in Russia, Finland, the
Volga, the Caucasus, and the Crimea.
February 15, 1899. General meeting. Thirty-two persons present.
Mr. Myron L. Fuller. Rapidity of sand-plain growth.
Prof. W. O. Crosby. Geology of the main dam and tunnel of
the Metropolitan water works, near Clinton, Mass.

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HENSHAW: REPORT OF SECRETARY AND LIBRARIAN. 19

February 17, 1899. Special meeting. Eighty-six persons present.
Prof. H. F. Osbom. Evolution of the Mammalia in North

America.

March 1, 1899. General meeting. Thirty-four persons present.

Dr. T. A. Jaggar, Jr. The intrusive rocks of the Black Hills.

March 15, 1899. General meeting. Twenty-eight persons present.

Mr. E. C. Jeffrey. Development and affinities of the genus

Equisetum.
Dr. C. R. Eastman. Some new North American fossil fishes.
April 5, 1899. General meeting. Sixty-three persons present.
Report of the Nominating committee.
Prof. C. E. Fay. The Alpine features of the Canadian

Rockies.
April 26, 1899. General meeting. One hundred and forty-four

persons present.
Dr. W. McM. Wood worth. Samoa and the Samoans.
Mr. G. M. Allen. Notes on the reptiles and amphibians of

Intervale, N. H. (By title.)
Dr. B. L. Robinson and Mr. J. M. Greenman. Contributions

from the Gray herbarium of Harvard university, no. 17.

(By title.)
Dr. Gary de N. Hough. Studies in Diptera Cyclorhapha.

(By title.)
Mr. F. N. Balch. List of marine MoUusca of Cold Spring

Harbor, Long Island, with descriptions of one new genus

and two new species of nudibranchs. (By title.)
Prof. G. E. Stone. The flora of Lake Quinsigamond. (By

title.)

PuBLirATIONS.

The following publications have been issued during the year : —

Localized stages in development in plants and animals, by Robert
T. Jackson. Memoirs, vol. 5, no. 4, 65 pp., 10 plates, 5 cuts.

The development, structure, and aftinities of the genus Equisetum,
by Edward C. Jeffrey. Memoirs, vol. 5, no. 5, 36 pp., 5 plates.

The genus Antennaria in New England, by Merrit L. Fernald.
Proceedings, vol. 28, no. 8, 13 pp.

Notes on a Carboniferous boulder train in eastern Massachusetts,
by Myron L. Fuller. Proceedings, vol. 28, no. 9, 14 pp., 1 cut.

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20 PROCEEDDfGS : BOSTON SOCIETY NATURAL HISTORY.

On the veins of the Wolffian bodies in the pig, by Charles S.
Minot. Proceedings, vol. 28, no. 10, 10 pp., 1 plate, 1 cut.

Proceedings of the annual meeting. May 4, 1898. Proceedings,
•vol. 28, no. 11,26 pp.

The odonate genus Macro themis and its allies, by Philip P.
Calvert. Proceedings, vol. 28, no. 12, 32 pp., 2 plates.

Some hydroids from Puget Sound, by Gary N. Calkins. Pro-
ceedings, vol. 28, no. 13, 35 pp., 6 plates.

North American wood frogs, by Reginald Heber Howe, Jr. Pro-
ceedings, vol. 28, no. 14, 6 pp.

Studies in the gold-bearing slates of Nova Scotia, by J. Edmund
Woodman. Proceedings, vol. 28, no. 15, 33 pp., 3 plates, 1 cut.

Moniloporidae, a new family of Palaeozoic corals, by Amadeus
W. Grabau. Proceedings, vol. 28, no. 16, 16 pp., 4 plates.

Library.

The additions to the library have

been as follows: —

8vo.

■tto.

Folio.

Total.

Volumes

278

309

Parts

2,208

373

2,582

Pamphlets

651

702

Maps

Total 3,137 430 71 3,638

The library contains 24,879 volumes, 1,378 incomplete (including
current) volumes, and 12,812 pamphlets.

By exchange, gift, or purchase we have added twenty-eight
serials: — Agricultural experiment stations of California, Con-
necticut, Delaware, Florida, Georgia, Illinois, Maine, Maryland,
Minnesota, New Mexico, Rhode Island, and South Carolina ; Bird
lore ; Blue Hill observatory bulletin ; Bulletin Cooper ornithologi-
cal club, Santa Clara ; Kansas university geological survey ; Lloyd
mycological museum, Cincinnati ; Mineral collector. New York ;
The museum, Albion, N. Y. ; Natural history association of Mira-
michi, Chatham, N. "B. ; Nature study in school. West Newton ;
Naturwissenschaftliche gesellschaft in Winterthur, Winterthur ;
Ottawa literary and scientific society, Ottawa ; Pasadena academy
of sciences, Pasadena; Queensland agricultural journal, Brisbane;

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HENSHAW: REPORT OF SECRETARY AND LIBRARIAN. 21

Rhodora, Boston ; Washington academy of science, Washington ;
Yorkshire naturalists' union, Leeds.

The Society now exchanges its publications with 432 scientific
institutions and periodicals.

One thousand and thirty-seven books have been borrowed by 115
persons ; 480 volumes have been borrowed for use in the building,
and the library has been consulted 450 times.

Three hundred and sixty-eight volumes have been bound in 824
covers ; 362 pamphlets have been bound.

Twenty-five volumes of the Verhandlungen zoologisch-botanische
gesellschaft, Wien, have been indexed ; current volumes of serials
previously indexed are indexed as received.

Taking advantage of the aid afforded by a generous though
anonymous patron, we have bound all the unbound books and
pamphlets on several alcoves in the back library; the alcoves
selected being those nearest the windows and thus the most exposed
to light and dust.

Walker Prizes.

The subjects selected by the Walker Prize Committee for 1899
were : —

1. Is there fundamental difference between 'equation division'
and * reduction division ' in the division of cells ?

2. The phenomena and laws of hybridization.

The only essay received, one on the relations between the hybrid
and parent forms of echinoid larvae, having been published in
Philosophical transactions of the royal society of London, could not
be considered by the Committee, as in all cases the memoirs are to
be based upon original and unpublished work.

It is greatly to be regretted that the annual awards of Walker
prizes are so frequently omitted, and it is suggested that the selec-
tion of subjects of broader scope would secure more general
competition.

The subjects for the award in May, 1900, are : —

1. Stratigraphy and correlation of the sedimentary formations of
any part of New England.

2. A study in palaeozoic stratigraphy and correlation.

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22 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

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ANNUAL MEETING. 23

The reports of the Trustees and of the Auditing Committee were
read and it was voted to accept and adopt the several reports.

The Society then proceeded to ballot for officers for 1899-
1900. Messrs. R. P. Bigelow and R. Hayward were appointed to
collect and count the votes. They reported the election of

PRESIDENT,

CHARLES SEDGWICK MINOT.

VICE-PRESIDENTS,

CHARLES p. BOWDITCH. HENRY W. HAYNES.

WILLIAM G. FARLOW.

CURATOR,

ALPHEUS HYATT.

SECRETARY,

samlt:l henshaw.

TREASURER,

EDWARD T. BOUVE.

LIBRARIAN,

SAMUEL HENSHAW.

councillor for three tears,

Miss Cora H. Clarke. George H. Parker.

Robert T. Jackson. A. Lawrence Rotch.

J. Arnold Lowell. William F. Whitney.

Edward 8. Morse. J. B. Woodworth.

councillor for one tear,
H. C. BuMPUs.

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24 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

OFFICERS FOR 1899-1900.

PRESIDENT,

CHARLES SEDGWICK MINOT.

VICE-PRESIDENTS,

CHARLES p. BOWDITCH. HENRY W. HAYNES.

WILLIAM G. FARLOW.

CURATOR,

' ALPHEUS HYATT.

SECRETARY,

SAMUEL HENSHAW.

TREASURER,

EDWARD T. BOUVfe.

LIBRARIAN,

SAMUEL HENSHAW.

COUNCILLORS FOR THREE YEARS,

Miss Cora H. Clarke.
Robert T. Jackson.
J. Arnold Lowell.
Edward S. Morse.

George H. Parker.
A. Lawrence Rotch.
William F. Whitney,
j. b. woodworth.

councillors for TWO YEARS,

Charles B. Davenport.
James H. Emerton.
William A. Jeffries.
George G. Kennedy.

Augustus Lowell.
Miss Susannah Minns.
Thomas A. Watson.
Samuel Wells.

S. L. Abbot.
William S. Bryant.
H. C. Bumpus.
William M. Davis.

COUNCILLORS FOR ONE YEAR,

Miss Catharine I. Ireland.
Benjamin Joy Jeffries.
N. T. Kidder.
Willla^m H. Niles.

George L. Goodale.

COUNCILLORS ez-officUs^

F. W. Putnam.
Samuel H. Sc udder.

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LIST OF MEMBERS.

HONORARY MEMBERS.

Alexander Agassiz,

Cambridge

Adolph Bastian,

Berlin.

John William Dawson,

Montreal.

Michael Foster,

Cambridge.

Ernst Haeckel,

Jena.

Joseph D. Hooker,

London.

Albert v. KoUiker,

Wtlrzburg.

Henri Lacaze-Duthiers,

Paris.

Edward B. Tylor,

London.

Kudolph Vlrchow,

Berlin.

WiUiam Allen,
James Anderson,
Francis Archer,
Francesco Ardissone,

Loring W. Bailey,
A. S. Baldwin,
Mariano Barcena,
A. Constantino Barry,
Charles E. Beecher,
Hermann Behr,
£. Tan Beneden,
William G. Binney,
Nathaniel H. Bishop,
William P. Blake,
George A. Boardman,
William H. Brewer,
William K. Brooks,
John Brown,

Giovanni Capellini,
Antonio del Castillo,
Ferdinand Cohn,
Guido Cora,

CORRESPONDING MEMBERS.

Boston.

Liverpool, Eng.
Liverpool, Eng.
Milan, Italy.

Fredericton, N. B.
Jacksonville, Fla.
Mexico, Mex.
Prairie du Sac, Wis.
New Haven, Ct.
San Francisco, Cal.
Li^ge, Belgium.
Burlington, N. J.
Lake George, N. Y.
New Haven, Ct.
Calais, Me.
New Haven, Ct.
Baltimore, Md.
New York, N. Y.

Bologna, Italy.
Mexico, Mex.
Breslau, Germany.
Rome, Italy.

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26 PKOCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

Elliott Coues,
John M. Coulter,
Hermann Credner,
EzraT. Cresson,
Josiah Curtis,

Henry Davis,
William Boyd Dawkins,
William Dean,
Anton Dohm,
Sanford B. Dole,
Henry E. Dresser,
Paul B. Du Chaillu,
W. T. T. Dyer,

Arthur M. Edwards,
William H. Edwards,
D. G. Elliot,
Sigmund Exner,

Roswell Field,
William H. Flower,
F. Fouqu6,

M. Ganin,
J. T. Gardner,
Albert Gaudry,
Archibald Geikie,
James Geikie,
Hans B. Geinitz,
Theodore N. Gill,
Augustus R. Grote,
Albert C. L. G. GUnther,
John T. Gulick,
L. H. Gulick,

Edwin Harrison,
James Hector,
Angelo Heilprin,
George Henslow,
Henry Y. Hind,
Charles H. Hitchcock,
John Hjaltalin,
W. J. Hoffman,
W. J. Holland,
Bernard A. Hoopes,
A. W. Howitt,
Oliver P. Hubbard,
Samuel Hubbard,

Wadiington, D. C.
Chicago, 111.
Leipsic, Germany.
Philadelphia, Pa.
Washmgton, D. C.

McGregor, Iowa.
Manchester, Eng.
Bangkok, Siam.
Naples, Italy?
Honolulu, H. I.
Kent, Eng.
New York, N. Y.
London, Eng.

Newark, N. J.
Coalburg, W. Va.
Chicago, 111.
Vienna, Austria.

GUI.

London, Eng.
Paris, France.

Nice, France.
Washington, D. C.
Paris, France.
Edinburgh, Scotland.
Edinburgh, Scotland.
Dresden, Germany.
Washington, D. C.
Hildesheim, Germany.
London, Eng.
Osaka, Japan.
Honolulu, Oahu.

St. Louis, Mo.
Wellington, N. Z.
Philadelphia, Pa.
London, Eng.
Windsor, N. S.
Hanover, N. H.
Rejkyavik, Iceland.
Reading, Pa.
Alleghany, Pa.
Philadelphia, Pa.
Melbourne, Vict.
Hanover, N. H.
San Francisco, Cal.

Digitized by VjOOQ IC

LIST OF MEMBERS.

Christopher Johnson,
David S. Jordan,

Clarence King,
John King,
Cornelius Kollock,
A. Kowalewsky,
Carl Kupffer,

Arnold Lang,

E. Ray Lankester,
Joseph Leconte,

R. von Lendenfeld,
J. Peter Lesley,
A. M. L6vy,

F. W. Lewis,
Franz Leydig,
Christian F. Ltitken,
Richard Lydekker,

Robert McLachlan,
E. J. Marey,
Paul Mayer,
Joseph B. Meader,
C. Hart Merriam,
Charles L. Metz,
Alphonse Milne-Edwards,
S. Wier Mitchell,
John Murray,

Francis P. Nash,
Alfred Newton,

Henry F. Osbom,

C. R. von Osten Sacken,

Emile Oustalet,

TTiomas F. Perley,
F6lix Plateau,
Edward B. Poulton,
John W. Powell,
Raphael Pumpelly,

Richard Rathbun,
Ferd von Richthofen,
Robert Ridgway,
Heinrich Rosenbtosch,

Henri de Saussure,
C. M. Scammon,

Baltimore, Md.
Lelstfid Stanford, Cal.

Washington, D. C.
Boone, Iowa.
Cheraw, S. C.
St. Petersburg, Russia.
Munich, Germany.

Zurich, Switzerland.
London, Eng.
Berkeley, Cal.
Czernowitz, Austria.
Milton.
Paris, France.
PhUadelphia, Pa.
Bonn, Germany.
Copenhagen, Denmark.
Harpenden, Eng.

London, Eng.
Paris, France.
Naples, Italy.
Stoneham.
Washington, D. C.
Madison ville, Ohio.
Paris; France.
Philadelphia, Pa.
Edinburgh, Scotland.

Geneva, N. Y.
Cambridge, Eng.

New York, N. Y.
Heidelberg, Germany.
Paris, France.

Portland, Me.
Li6ge, Belgium.
Oxford, Eng.
Washington, D. C.
Newport, R. I.

Washington, D. C.
Bonn, Germany.
Washington, D. C.
Heidelberg, Germany.

Geneva, Switzerland.
Washington, D. C.

Digitized by VjOOQ IC

28 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

Philip L. Sclater,
A. R. C. Selwyn,
William Sharswobd,
Hamilton L. Smith,
Hermami Snellen,

Armand Thieleus,
Tamerlan Thorell,
William Trelease,
Gustav Tschermak,

Philip R. Uhler,

Addison E. Verrill,
Sydney H. Vines,

W. Waagen,

Henry A. Ward,

R. A. Ward,

Carl Wedt,

August Weismann,

George M. Wheeler,

WUliam T. White,

R. P. Whitfield,

Robert E. E. Wiedersheim,

Burt G. Wilder,

C. S. Wilkinson,

Edmund B. Wilson,

Henry Woodward,

J. J. Woodward,

Ferdinand Zirkel,
Carl A. Zittel,

London, Eng.
Ottawa, Can.
Philadelphia, Pa.
Geneva, N. Y.
Utrecht, Holland.

Tirlemont, Belgium.
Montpellier, France.
St. Louis, Mo.
Vienna, Austria.

Baltimore, Md.

New Haven, Ct.
Oxford, Eng.

Vienna, Austria.
Rochester, N. Y.
Troy, N. Y.
Vienna, Austria.
Freiburg, Germany.
Washington, D. C.
New York, N. Y.
New York, N. Y.
Freiburg, Germany.
Ithaca, N. Y.
Sydney, N. S. W.
New York, N. Y.
London, Eng.
Washington, D. C.

Leipsic, Germany.
Munich, Germany.

CORPORATE MEMBERS.

Samuel L. Abbot, M. D.,
John E. Alden,
Jane Alexander,
Henry F. Allen,
Joel A. Allen,
Edward P. Allis, Jr.,
Arthur Amory,
Robert Amory, M. D.,
S. Reed Anthony,
Nathan Appleton,
William S. Appleton,
Anuetta F. Armes,
Edward P. Austin,

00 Mt. Vernon St.
Newton.

91 Mt. Vernon St.
Jamaica Plain.
Absent.

Milwaukee, Wis.
133 Marlborough St.
279 Beacon St.
63 State St.
Somerset Club.
462 Beacon St.
Dorchester.
Absent.

Digitized by VjOOQ IC

LIST OF MEMBERS.

Lucas Baker,
Francis N. Balch,
Edward E. Bancroft, M. D.,
Edward A. Bangs,
Outram Bangs,
James M. Barnard,
Walter B. Barrows,
George H. Barton,
Charles F. Batchelder,
George W. Beaman,
Mrs. George W. Beaman,
Henry B. Bigelow,
Joseph S. Bigelow, Jr.,
Robert P. Bigelow,
William S. Bigelow, M. D.,
aarence J. Blake, M. D.,
Francis Blake,
James H. Blake,
Joseph W. Blankinship,
Albert N. Blodgett, M. D.,
>Ir8. Alice L. Boardman,
Elizabeth D. Boardman,
Edward T. Bouv6,
Charles P. Bowditch,
Frederic C. Bowditch,
Henry P. Bowditch, M. D.,
Mrs. Ella F. Boyd,
Arthur C. Boy den,
Francis W. Brewer,
Mrs. Thomas M. Brewer,
Willard S. Brewer,
Edwin T. Brewster,
William Brewster,
Albert P. Brigham,
Henry Brooks,
Shepherd Brooks,
J. Frank Brown,
Edward I. Browne,
Elizabeth B. Bryant,
John Bryant, M. D.,
William S. Bryant, M. D.,
Charles Bullard,
Williani N. Bullard, M. D.,
Hermon C. Biunpus,

Absent.

Jamaica Plain.

Wellesley.

240 Beacon St.

Milton.

Absent.

Mass. Inst, of Technology.

Cambridge.

Absent.

Cambridge.

261 Commonwealth Ave.

251 Commonwealth Ave.

Mass.. Inst, of Technology.

60 Beacon St.

226 Marlborough St.
Auburndale.
Cambridge.
Bozeman, Mont.

61 Massachusetts Ave.
Absent.

416 Marlborough St.

436 Marlborough St.

28 State St.

Brookline.

Harvard Medical School.

Hyde Park.

Bridgewater.

Hiugham.

233 Beacon St.

Hingham.

Andover.

Cambridge.

Absent.

Lincoln.

02 Beacon St.

89 State St.

63 State St.

AUston.

Cohasset.

63 State St.

Cambridge.

89 Marlborough St.

Providence, R. I.

Arthur T. Cabot, M. D.,
Edward C. Cabot,
Louis Cabot,

1 Marlborough St.

Brookline.

Digitized by VjOOQ IC

30 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

William B. Cabot,

Mrs. Gar>' N. Calkins,

Alvin Carl,

Charles T. Carruth,

Mrs. J. B. Case,

Arthur P. Chadbourne, M. D.,

Montage Chamberlain,

Walter G. Chase,

Francis S. Child,

Henr>' L. Clapp,

Mabel D. Clapp,

John S. Clark,

T. W. B. Clark,

Cora H. Clarke,

Collier Cobb,

Edward W. Ctxlman,

Helen Collamore,

Frank S. Collins,

Grace E. Cooley,

Algernon Coolidge, M. D.,

T. Jefferson Coolidge, Jr.,

Mrs. Elizabeth R. Cormier,

Charles B. Cor>',

Charles U. Cotting,

Sarah H. Crocker,

William O. Crosby,

Charles R. Cross,

Clara E. Cmnmiugs,

Chestnut Hill.

New York, N. Y.

Jamaica Plain.

84 India St.

468 Beacon St.

226 Marlborough St.

Cambridge.

Brookline.

Absent.

Roxbury.

9 Mas.sachusetts Ave.

046 Washington St.

326 Atlantic Ave.

01 Mt. Vernon St.

Chapel Hill, N.C.

63 Marlborough St.

317 Commonwealth Ave.

Maiden.

Wellesley.

81 Marlborough St.

184 Beacon St.

Charlestown.

100 Boylston St.

249 Commonwealth Ave.

319 Commonwealth Ave.

Mass. Inst, of Technology.

Wellesley.

Ulric Dahlgren,
William H. Dall,
Reginald A. Daly,
Ada Daiia,
John Dane,
F. Graef Darlington,
Charles B. Davenport,
Andrew McF. Davis,
James C. Davis,
Simon Davis,
William M. Davis,
Henry G. Denny,
Franklin Dexter, M. D.,
F. Gordon Dexter,
Gordon Dexter,
Joseph S. Diller,
George Dimmock,
Richard E. Dodge,
Charles C. Doe,

Princeton, N. J.

Absent.

Newton.

29 Marlborough St.

Absent.

Cambridge.

70 Kilby St.

277 Beacon St.

Cambridge.

08 Devonshire St.

148 Marlborough St.

65 Beacon St.

66 Beacon St.
Absent.
Spriugtield.
Absent.

South Newbur>', Vt.

Digitized by VjOOQ IC

LIST OF MEMBERS.

Thomas Doliber,
Jonathan Dorr,
George A. Dorsey,
Richard S. Dow,
Sara A. Downs,
Thomas Dwight, M. D.,
Harrison G. Dyar,

Raymond B Earle,
Charles R. Eastman,
Charles W. Eliot,
Mrs. J. W. Elliot.
Mary L. Ells,
James H. Emerton,
William Endicott,
Harold C. Ernst, M. D.,
James F. Estes,

William G. Farlow, M. D.,
Mrs. Eva D. Farquhar,
Charles E. Faxon,
Heiirj' H. Fay,
Joseph S. Fay.
Charles S. Fellows,
Cliarles H. Fernald,
J. Walter Fewkes,
William L. W. Field,
Frank S. Fiske,
George W. Fitz, M. D.,
Augustus Flagg,
Charles F. Folsom, M. D.,
Justus W. Folsom,
Eugene N. Foss,
John Foster,
William Foster,
Harriet .E. Freeman,
Nathaniel S. French,
Sophia W. French,
Charles Fr>',
Myron L. Fuller,
Sarah S. Fuller,

Thomas Gaffield,
Charles W. Galloupe,
T. W. Galloway,
William F. Ganong,
Edward G. Gardiner,
John L. Gardner, Jr.,

Brookline.
27 School St.
Chicago, 111.
27 State St.
68 Berkeley St.
236 Beacon St.
Washington, D. C.

Newton.

Cambridge.

124 Beacim St.

Cambridge.

Clarendon St.

33 Summer St.

Hanard Medical School.

Absent.

Cambridge.

Roxbury.

Jamaica Plain.

416 Beacon St.

Id9 Commonwealth Ave.

Fairbanks, Fla.

Amherst.

Absent.

Milton.

U. S. Court House.

Cambridge.

274 Clarendon St.

16 Marlborough St.

Cambridge.

Jamaica Plain.

113 Broad St.

Absent.

384 Commonwealth Ave.

West R«>xbury.

Wollaston.

21 Commonwealth Ave.

Mass. Inst, of Technology.

Absent.

64 Allen St.

46 Broad St.

Marshall, Mo.

Absent.

131 Mt. Vernon St.

61 Commonwealth Ave.

Digitized by VjOOQ IC

32 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

George M. Garland, M. D.,

George W. Gay, M. D.,

B. W. Gilbert,

George L. Goodale, M. D.,

Harold B. Groodrich,

^Irs. Mary T. Gorhain,

Amadeus W. Grabau,

Francis C. Gray,

John C. Gray,

Robert W. Greenleaf, M.D.,

Mrs. C. W. Greenough,

Charles P. Greenough,

H. S. Greenough,

William D. Grier,

Leon S. Griswold,

227 Newbury St.

665 Boylston St.

202 Purchase St.

Cambridge.

72 Westland Ave.

108 Marlborough St.

Cambridge.

7 Mt. Vernon Place.

176 Beacon St.

561 B»)ylston St.

Jamaica Plain.

39 Court St.

Absent.

20 Kilby St.

Dorchester.

Minna B. Hall,
Robert W. Hall,
Susan M. Hallowell,
Mrs. M. L. Hammatt,
Ida S. Hammerle,
Edward D. Harris,
T. W. Harris,
Francis R. Hart,
Franklin Haven,
Gustavus Hay, M.D.,
Ellen Hayes,
Henry W. Haynes,
Roland Hayward,
Charles E. Hellier,
Augustus Hemenway,
Mrs. Augustus Hemenway,
Joseph P. B. Henshaw,
Samuel Henshaw,
Warren W. Herman,
Francis H. Herrick,
Ella J. Hill,
Harriet A. Hill,
Hibbert W. Hill, M. D.,
Mary H. Hinckley,
John Hobbs,
Walter E. Hobbs,
John Hogg,
Frederick S. HoUis,
John Homans, M. D.,
Robert C. Hooper,
Mrs. S. E. Hooper,
Samuel A. Hopkins, '.M. D.,

Brookline.

Cambridge.

Wellesley.

Hyde Park.

Roxbury.

Absent.

Milton.

07 Mt. Vernon St.

388 Marlborough St.

Wellesley.

289 Beacon St.

346 Marlboroui?h;St.

57 Etiuitable Bldg.

273 Clarendon St.

77 Newbury St.

Cambridge.

P. O. Box 1848.

Cleveland, Ohio.

223 Newbury St.

Belmont.

72 Pinckney St.

Mattapau.

99 St. Botolph St.

Stony brook.

280 Commonwealth Ave.

Newton Highlands.

164 Beacon St.

448 Beacon St.

Cambridge.

235 Marlborough St.

Digitized by VjOOQ IC

LIST OF MEMBERS.

Gany de N. Hough, M.D.,
Theodore Hou^,
Helen Hubbard,
John G. Hubbard,
L. L. Hubbard,
John E. Hudson,
Henry S. Himnewett,
VVillard P. Huimewell,
Alpheus Hyatt,

Charles E. Inches, M. D.,
Catherine I. Ireland,

John G. Jack,
John C. Jackson,
Robert T. Jackson,
William D. Jackson,
Thomas A. Ja^jgar, Jr.,
B. Joy Jeffries, M. D.,
William A. Jeffries,
Charles W. Jenks,
Charles F. Jenney,
Isabel L. Johnson,
Samuel Johnson,
Marian H. Judd,

Charles S. Kendall,
Mrs. Caroline A. Kennard,
Getjrge G. Kennedy, M. D.,
Harris Kennedy, M. I).,
Nathaniel T. Kidder,
John S. Kingsley,

New Bedford.

Mass. Inst, of Technology.

(^harlestown, N. H.

Brookline.

Absent.

126 Milk St.

9 Park St.

261 Commonwealth Ave.

Cambridge.

380 Beacon St.
Cambridge,

Jamaica Plain.

Absent.

3.S Gloucester St.

Bridgewater.

Cambridge.

16 Chestnut St.

126 Beaccm St.

Bedford.

Hyde Park.

467 Massachusetts Ave.

7 Commonwealth Ave.

186 Conmionwealth Ave.

91 Federal St.

Brookline.

Readville.

Milton.

Tufts College.

F. D. Lambert,
Alfred C. Lane,
Amory A. Lawrence,
William Lawrence,
Geora^e W. Ixie,
David F. Lincoln, M. D.,
James L. Little,
William R. Livennore,
William C. Loring,
Augustus Ivowell,
Charles Lowell,
James Arnold Lowell,
Mrs. Louisa F. Lowery,
Arthur T. Lyman,
Charles P. Lyman, M. D.,

Auburn, Me.

Houghton, Mich,

69 (\)nunonwealth Ave.

122 (Commonwealth Ave.

Brookline.

73 Pinckney St.

Brookline.

P. O. Box 168.

2 Gloucester St.

63 State St.

149 Beacon St.

297 Beacon St.

AbstMit.

P. O. Box 1717.

60 Village St.

Digitized by VjOOQ IC

34 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

George H. Mackay,
W. Duncan MoKim,
J. Playfair McMurrich,
W. D. McPherson,
B. Pickman Mann,
Warren H. Manning,
Edward L. Mark,
Mrs. W. H. K. Marrs,
Vernon F. Marsters,
Asa E. Mattice,
F. W. G. May,
Alfred G. Mayer,
Charles J. Maynard,
James Means,
James C. Merrill, M. D.,
Selah Merrill,
Gerrit S. Miller, Jr.,
Susannah Minns,
Charles S. Minot,
Laurence Minot,
William Minot,
George Mixter,
William J. Moenkhaus,
Henry L. Moody,
Alexander Moore,
Margaret W. Morley,
Albro D. Morrill,
Albert P. Morse,
Edward S. Morse,
Elisha W. Morse,
John Murdoch,
Albert L. Murdock,

Nathaniel C. Nash,
Herbert V. Neal,
Frederick H. Newell,
Mrs. Edith J. Nichols,
Sereno D. Nickerson,
William H. NUes,
Grenville H. Norcross,
Edward E. Norton,
William E. Norton,

John Ome, Jr.

Alpheus S. Packard, M. D.,
George H. Parker,
Edith A- Parkhurst,
George L. Parmelee,

218 Commonwealth Ave.
Portsmouth, N. H.
Absent.

South Framingham.

Absent

Brookline.

Cambridge.

Absent.

Bloomington, Ind.

Concord, Mich.

Dorchester.

Cambridge.

West Newton.

106 Beacon St.

Absent,

Absent.

Washington, D. C.

14 Louisburg Sq.
Harvard Medical School.
24 Marlborough St.

39 Court St.

219 Beacon St.
Williamstown.
Absent.

3 School St

28 St James Ave.

Clinton, N. Y.

South Natick.

Salem.

Jamaica Plain.

Roxbury..

15 Causeway St.

Cambridge.

Absent.

Absent

294 Marlborough St.

Masonic Temple,

Cambridge.

9 Commonwealth Ave.

419 Washington St

Absent.

Cambridge.

Providence, R. L
Cambridge.
Somerville.
Absent.

Digitized by VjOOQ IC

LIST OF MEMBERS.

William Patten,
Francis H. Peabody,
James £. Peabody,
Edward C. Perkins,
William H. Phelps,
John C. Phillips,
Dudley L. Pickman,
David Pingree.
Julia B. Piatt,
William G. Preston,
Frances C. Prince,
Loring W. Puffer,
Charles P. Putnam, M. D.,
Frederick W. Putnam,
James J. Putnam, M. D.,

Hanover, N. H.

113 Devonshire St.

Absent.

706 Sears Bldg.

Absent.

299 Berkeley St.

98 Beacon St.

Salem.

Absent.

186 Devonshire St.

17 Joy St.

Brockton.

63 Marlborough St.

Cambridge.

106 Marlborough.

Motte A. Read,

Mrs. William H. Reed,

John P. Reynolds, M. D.,

Stephen H. Rhodes,

Mrs. Ellen H. Richards,

George H. Richards,

Harriet E. Richards,

Robert H. Richards,

William L. Richardson, M. D.,

Everett W. Ricker,

William Z. Ripley,

Thomas P. Ritchie,

Benjamin L. Robinson,

Alfred P. Rockwell,

Mrs. William B. Rogers,

A. Lawrence Rotch,

WUUam H. Ruddick, M. D.,

Mrs. T. E. Ruggles,

John D. Runkle,

Frederick W. Russell, M. D.,
«

William E. Safford,
Lilian V. Sampson,
Charles S. Sargent,
Dudley A. Sargent, M. D.,
Frederick Le R. Sargent,
Mrs. Marian E. Y. Saville,
Marshall H. Saville,
Henry Sayles,

Alfred L. T. Schaper, M. D.,
Barthold Schlesinger,
Samuel H. Scudder,

Absent.

37 Commonwealth Ave.

416 Marlborough St.

641 Commonwealth Ave.

Jamaica Plain.

14 Chestnut St.

Boston.

Jamaica Plain.

225 Commonwealth Ave.

City Hall.

Mass. Inst, of Technology.

Newton Highlands,

Cambridge.

281 Beacon St.

117 Marlborough St.

63 State St.

602 E. Broadway.

Milton.

Mass. Inst, of Technology.

Winchendon.

Absent.

Germantown, Pa.

Brookline.

Cambridge.

Absent.

Waban.

Absent.

Somerset Club.

Harvard Medical School.

131 Devonshire St.

Cambridge.

Digitized by VjOOQ IC

PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

John H. Sears,

Salem.

Mrs. Mary L. Seavey,

Brookline.

William T. Sedgwick,

Mass. Inst, of Tt*chnology

Nathaniel S. Shaler,

Cambridge.

J. C. Sharp, Jr.,

64 Commonwealth Ave.

Stephen P. Sharpies,

13 Broad St.

Mrs. J. M. Sheldon,

108 Mt. Vernon St.

Augustine Shurtleff, M. D.,

Brookline.

A. D. Sinclair, M. D.,

35 Newbury St.

Charles C. Smith,

286 Marlborough St.

Frederic F. Smitli,

Spriiigfield.

Caroline G. Soule,

Brookline.

Edmund D. Spear, M. D.,

20 Mt. Vernon SU

A. W. Spencer,

31 State St.

Charles J. Sprague,

380 Marlborough St.

Frank F. Stanley,

108 Summer St.

George E. Stt)ne,

Amherst.

Charles S. Street,

Absent.

Mrs. Charles P. Strong,

Cambridge.

William C. Sturgis,

New Haven, Ccmn.

John 0. Sumner,

Ab.M'nt.

Charles W. Swan, M. D.,

Brookline.

J. Brooks Taft,

28 Peniberton S(i.

Ralph S. Tarr,

Absent.

Levi L. Thaxter,

13 Tremont St.

Roland Thaxter,

Cainbrldm'.

John E. Thayer,

Lancaster.

Mary F. Thompson,

413 Shawmut Ave.

Augustus L. Thorndikc,

722 Treinont Bldg.

Townsend W. Thorndikc,

22 Newbury St.

G. Francis Topliff,

48 Congi-ess St.

Mrs. Helen M. Tower,

Cainbridgeport.

Samuel F. Tower,

English High School.

William L. Tower,

Cambridge.

W. Porter Truesdell,

12 South St.

Frederick Tuckerman, M. D.,

Andierst.

William Tudor,

Absent.

Warren Upham,

Absent.

J. F. Urie, M. D.,

Absent.

Balfour H. Van Vleck,

Boston Soc. of Nat. Hist.

T. Wayland Vaughan,

Absent.

M. Edward Wadsworth,

Absent.

Oliver F. Wadsworth, M. D.,

520 Beacon St.

Robert Wainwright,

Absent.

Digitized by VjOOQ IC

LIST OF MEMBERS.

Frederick C. Waite,
Mary L. Ware,
Joseph W. Warren, M. D.,
Mrs. Elizabeth S. Wat«on,
Thomas A. Watson,
Mary M. Webster,
Clarence M. Weed,
Andrew G. Weeks,
Andrew G. Weeks, Jr.,
Charles G. Weld, M. D.,
Samuel M. Weld,
Samuel Wells,
William P. Wesselhoeft,
Arthur W. Weysse,
Mrs. Katharine K. Wheeler,
Charles T. White,
James C. White, M. D.,
Charles O. Whitman,
Henry M. Whitney,
Solon F. Whitney,
William F. Whitney, M. D.,
W. H. Whittemore,
George Wigglesworth,
Thomas Wigglesworth,
Mary A. Willcox,
Emile F. Williams,
Henry V. Wilson,
WiUiam F. WUson,
ClifU>n E. Wing, M. D.,
Guy M. Winslow,
Roger Wolcott,
John E. Wolff,
Edward S. Womi, M. D.,
Elvira Wood,
J. Edmund Woodman,
Frederick A. Woods, M. D.,
Jay B. Woodworth,
William McM. Woodworth,
G. Frederick Wright,

New York, N. Y.
41 Brimmer St.
Absent.
Weymouth.
Weymouth.
232 Newbury St.
Durham, N. H.
400 Beac(m St.
360 Washington St.
6 Commonwealth Ave.
North Chathaju.
46 Commonwealth Ave.
M. D., 176 Commonwealth Ave.

Mass. Inst, of Tech noli ►gy.
Roxbury.

213 Connnonwealth Ave.
250 Marlborough St.
Chicago, III.

107 (^)mmonwealth Ave.
Wjit^rtown.
228 Mariborough St.
Milfnrd, N. H.
53 State St.
36 Hawhy St.
WelU'Kh.y.

352 MiiKsachusctts Ave.
Chapt'l Hill, N. C.
Abst^nt.
Hoxbuiy.
Auburndale.
173 Commonwealth Ave.
Cambridge.

Harvanl Medical S<*hool.
Mjuss. Inst, of Technology,
(^and^ridgf.
Brookline.
Cambridge.
C'ambridge.
ObeHin, Ohio.

E. Bentley Young,

Francis Zirngiebel,

104 Apploton St.
Roxbury.

Digitized by VjOOQ IC

38 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

PATRONS.

Loring W. Bailey. Nathan Matthews.

W. W. BaUey. John J. May.

James M. Barnard. Samuel May.
Miss Emellne Binney.

Mrs. Mary G. P. Binney. Henry Sayles.

Mrs. G. H. vShaw.

George R. Carter. Charles J. Sprague.

Henry Cross. W. W. Stone.

Thomas Wigglesworth.
C. W. Galloupe. Edward Wyman.

BY-LAWS.

Section 1. Membebs.

1. The Society shall consist of Corporate Members, Corre-
sponding Members, Honorary Members, and Patrons.

2. Members shall be elected by the Council only upon the
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8. Corresponding and Honorary Members may be chosen from
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when the Council shall so order.

5. A Member may be expelled from the Society upon the
recommendation of the Council, by a vote of three fourths of the

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BY-LAWS. 39

Members present at a meeting of the Society after notice of such
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Society, a sum not less than five hundred dollars shall become a
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Section 2. Admission Fee and Assessments.

1. Every person elected a Corporate Member, before obtaining^
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4() PROC EEDINGS : BOSTON SOCIETY NATURAL HISTORY.

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All Presidents and Vice-Presidents prior to January 7, 1891, are
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All officers shall l>e chosen by ballot. Of the twenty-four
Councillors eight shall be chosen at each Annual Meeting to serve
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and of the Council.

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his absence in the order of seniority in office.

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5. The Secretary shall record the proceedings of the Society,
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notice of all meetings, and inform officers of all matters which
occur at any meeting requiring their action.

6. The Librarian shall have charge of the library and shall
observe and enforce such regulations as the Council may make for
its use.

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BY-LAWS. 41

7. The Treasurer shall receive all moneys due or payable to the
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approved by the Executive Committee ; and shall keep a correct
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1. The Council shall have full power to act for the interests of
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the Curator, the Secretary, and the Treasurer.

2. The Annual Meeting of the Council shall be held on the
second Wednesday in May, at which meeting a Board of three
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Society in trust with power to sell and to reinvest according to
their judgment. The following Standing Committees shall also be
elected ; viz. : an Executive Committee of five, in whom the affairs
of the Council shall be vested so far as it shall determine by its
votes and rules; a Membership Committee of five to recommend
to the Council persons for election as Members; a Library and
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issue and exchange of the publications of the Society; a Walker
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accounts of the Treasurer. These Committees and Boards need
not be chosen from the Council, and they shall hold office until
their successors have qualified.

Special committees also may be appointed if needed.

3. The Council may fill the vacancies occurring among the
twenty- four Councillors until the next Annual Meeting of the
Society.

Section 6. Meetings.

1. The Annual Meeting for the election of officers and for
other general purposes shall be held on the first Wednesdajr in

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42 PROCEEDINGS : BOSTON SO(^IETY NATURAL JIISTORY.

May. At this meeting the following reports sliall be presented :
bj the Curator uj)on tlie condition and progress of tlie museum,
the lectures which he has superintended, and any other matters
of general interest; by the Secretary upon the publications,
meetings, and the lectures which he has superintended ; by the
Librarian upon the library; by the Treasurer upon the receipts
and expenditures for the year; and by the Trustees upon the
tinancial condition of the Society.

2. General meetings of the Society shall be held on the first
and third Wednesdays of every month from November to May
inclusive ; and at such other times as the Council may determine.

When any meeting provided for by these By-Laws falls upon a
legal holiday, said meeting shall be held on the Wednesday follow-
ing such holiday.

3. At any meeting business not provided for by these By-Laws
may be transacted, provided notice of such ])roposed business has
been mailed postpaid at least four days before the meeting to the
address furnished by each Corporate ^Member.

4. Eleven Corporate Members shall form a quorum for business.

5. The order of proceeding at the (General Meetings shall be

1. Approving the record of preceding meeting.

2. Business.

3. Scientific communications and their discussion.

4. Adjournment.

Section 7. Libkaky.

1. Members of the Society shall have access to and may take
books from the library under the direction of the Library and
Publishing Committee, which may by special vote extend the use
of the books to others than Members, specifying the conditions
under which books may be taken, or may restrict their use even by
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demand it.

2. The rules and regulations for the use of the library shall be
posted in the library room, and a digest of them aftixed to each
volume.

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BY-LAWS. 43

Section 8. Museu3i.

1. The Executive Committee shall determine the conditions of
admission to the museum.

2. Specimens may be removed from the museum only by leave
of the Curator who shall take a receipt of the same and be respon-
sible for their return in good order.

Section 9. Sections.

1. The Council may authorize the formation of Sections of the
Society on the written ai)plication of ten Members.

2. Meetings may be held at the pleasure of the Section.

3. The requirements of membership in a Section shall be : —

Membership in the Society.

Written nomination by two Members of the Section at

one of its regular meetings.
Election by a three fourths vote of the Members present

at a subsequent meeting of the Section.

4. The records of meetings of Sections shall be entered in
chronological order upon the book containing the records of the
ordinary meetings of the Society.

6. Such report of each meeting as may be judged by the Library
and Publishing Committee suitable for publication in the Proceed-
ings or Memoirs of the Society, shall be announced by the Secretary
at the next regular meeting of the Society.

6. Each Section shall have the right to make additional regula-
tions concerning its organization, subject to the approval of the
Council.

Section 10. Change of By-Laws.

1. The By-Laws of the Society may be altered or amended by a
two thirds vote of the Corporate Members present at any meeting,
provided a notice of such intended change shall have been given at
a preceding meeting and shall have been mailed postpaid to the
address furnished by each Corporate Member, at least four days
before final action.

Printed, June, 1899.

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Proceedings of the Boston Society of Natural History.

Vol. 29, No. 2,

p. 45-61.

VARIATION AND SEXUAL SELECTION IN MAN.

Bir Edwin Tennky Brewster.

BOSTON:

PRINTED^ FOR THE SOCIETY.

''July, 1899.

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No. 2. — Variation and Sexual Selection in Man,

By Edwin Tenxet Brewster, Andover, Mass.

The proverbial *inch on a man's nose' suggests the rather
obvious fact that certain parts of the human body determine the
personal appearance far more than do other parts. These portions,
which may be no larger or more useful than others, I shall (for
want of a better term) call conspicuous. This paper attempts to
show that there is a relation between the conspicuousness of any
part of the body, and its variability as measured by the coefficient
of variability of- its dimensions. ( Cy. Pearson, '96, p. 265-277 ;
Brewster, '97, p. 269-273. I here follow Professor Pearson in
multiplying all coefficients by 100.) The data of the investigation
are given in such of the appended tables as are designated by
letters, while comparisons and synopses are given in the numbered
Ubles.

I shall first offer evidence to prove that conspicuous dimensions
tend to be more variable than other dimensions. Table A, the first
to be considered, gives the ^ coefficient of variability ' of ten bone
measurements for New England Indians of both sexes. From this
it appears that the several dimensions of the head and face have
coefficients of variability which may be as low as 2.5 or as high as 5.
In other words, some dimensions, when measured by this method,
are twice as variable as others. Now it is evident that a person's
appearance is determined by the dimensions of the face rather than
by those of the head ; that it is, as a whole, the face rather than the
head, which is noticed and remembered ; which is, in short, con-
spicuous. This, of course, does not mean that every feature of
the face is more conspicuous than any feature of the head. The
width of the jaw, for example, since it is partly masked by the
cheeks, affects the appearance decidedly less than the size and shape
of the forehead. But the head, as a whole, is less conspicuous than
the face as a whole. If, then, conspicuousness is correlated with
variability, the dimensions of the face should be more variable than
those of the head, and the mean of the six coefficients of the face
dimensions should be greater than the mean of the four coefficients
of the head dimensions. A glance at Table A shows that this rela-

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46 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

tion holds, for the coefficients of the head are obviously smaller than
those of the face.

Kot only may we say that the face is more conspicuous than the
head, but we may go farther and say that the size and shape of the
nose affect the appearance more than any other feature of the face.
These three propositions should then be true : —

(1) The nose should be more variable than the head.

(2) The face without the nose should be more variable than the
head.

(3) The nose should be more variable than the rest of the face.
In the first colunm of Table 1 b given, for both sexes, the mean

of the coefficients of variability of the four head dimensions. Column
4 of the same table gives the mean of the coefficients of the four
dimensions of the face, and column 5 the mean of the coefficients of
the two nose dimensions. From these figiires it appears, that (1)
is true for both sexes ; that (2) b also true for both sexes ; while
(3) , though true for the females, is not true for males. Under col-
umns 3 and 6 are given the scores for these two tests. Beneath the
sign -f are given the cases of correlation between conspicuousness
and variability ; under the sign — the cases in which correlation is
wanting. The two tests give a score of 7 to 1 in favor of correla-
tion.

The data of Table A may be made to furnish yet another test of
this relation. A moment's consideration will show that we nearly
always visualize other persons in full face view rather than in pro-
file, and that we think of other men and races as they look when
seen face to face. It follows, then, that transverse diameters of the
head and face, which determine the full face aspect, are, in general,
more conspicuous than the dorso-ventral dimensions, which are seen
more clearly in profile ; and that vertical dimensions, which affect
both aspects, are more conspicuous than either. Here, again, I do
not imply that every transverse dimension affects the appearance
more than any dorso-ventral dimension, but only that this statement
is true in general. We shall expect to find, then, (1) that the mean
coefficient of the three dorso-ventral dimensions is less than the
mean of the four transverse dimensions, and (2) less than the mean
of the four vertical dimensions, and (3) that the mean of the trans-
verse dimensions is less than the mean of the vertical dimensions.
Columns 7, 8, 9 give these means, and show that all three state-
ments are true for women, but only the third is true for men. The

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BREWSTER : VARIATION AND SEXUAL SELECTION IN MAN. 47

score for this test, recorded as before, is given in column 10, and is
4-2 for both sexes. Column 11 gives the total score for the three
tests which have been applied to the data of Table A. Altogether,
in eleven cases out of fourteen, conspicuousness is associated with
Tariability.

In TM» B are given the coefficients of various skull and face
dimensions of Venezuelans, and in Table 2 I have brought together
averages corresponding to those of the New England Indians given
in Table 1. If the same comparisons are made among these aver-
ages as were made among tho0e of Table 1, in every case the more
conspicuous dimensiojis are seen to be the more variable.

The first ten columns of Table 3 repeat^ for the data of Table C,
the tests already applied to the data of Tables A and B. Column 3
shows seven correct cases out of a possible nine ; column 6, twenty-
one out' of twenty-seven; column 10 shows nineteen out of twenty-
seven ; a total of 47 out of 63. All the tests thus far applied give a
total of 86 to 19, or something more than 80 per cent.

Suppose now, in Table C, I consider the face measurements alone,
and divide them into two groups, one of which shall contain
the coefficients of all the more conspicuous dimensions, and the
other the coefficients of the less conspicuous. Concerning the con-
spicuousness of some of these dimensions I am unable to form an
opinion. But I think I am tolerably safe in claiming that the dis.
tance from the ear to the nose root, the distance between the
temples (upper face breadth) and the distance between the angles
of the jaw (lower face breadth) are less conspicuous than the under
jaw length, cheek breadth, and mouth breadth. I may state at this
point that I made the division first, and computed the mean after-
ward. Column 11 gives the mean coefficient of the three less con-
spicuous dimensions, and column 12 the mean for the more conspic-
uous. In each of the nine cases the relative magnitude of the
coefficient is what would be predicted.

I may go even farther, and compare four conspicuous with four
inconspicuous dimensions, by including with the conspicuous dimen-
sions already taken, the distance from ear to chin, and with the
inconspicuous dimensions, the distance between the inner angles of
the eye (nose root breadth). These two dimensions are not so
clearly assignable to their respective classes as the six dimensions
first compared. Nevertheless, the distance from ear to chin meas-
ures the protnision of the lower jaw, one of the most conspicuous

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48 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

features of the lower part of the face ; on the other band, the nose
root breadth, though by no means inconspicuous, is, I think,
obscured by the more prominent features near it. At any rate,
these are the only two face dimensions remaining, which in any-
wise lend themselves to this classification. Columns 13 and 14,
incomplete through lack of data, show in turn complete correlation.
There is, however, one head dimension which is eminently con-
spicuous, the forehead height. This, in column 16, 1 have included
with the dimensions of column 14; and in column 15, with the
dimensions of column 13, 1 have included the breadth between the
ears. I have chosen this last dimension because it is apparently
the most variable of the remaining head dimensions, and, in conse-
quence, least favorable to my case. Here, too, in comparing four
conspicuous dimensions of the face and one of the head with four
inconspicuous dimensions of the face and one of the head,* I find
in every instance that the more conspicuous dimensions are the
more variable.

It seems, at first sight, an easy matter to apply this last test to
the data of Tables A, B, and D. But in the case of Table A the
number of measurements is too few, and in Tables B and D I have
found it impossible to assign the data to the proper groups.

Finally, in column 18, are given the mean coefiicients of eight
body measurements ; these should evidently be less than the coeffi-
cients of the more conspicuous face dimensions. So indeed they
are, except in the case of the Chinese ; and even here, as in every
instance, the face including the nose is more variable than the body.
Table D is like A and B, and Table 4 like 1 and 2, and need no
comment; they give twelve good cases out of fourteen.

I have now made 142 comparisons between the variability of
various dimensions classified according to their importance for per-
sonal appearance; and in 120 cases — more than 84% — the more
conspicuous dimensions are the more variable. This per cent would
doubtless be greater, if the coefiicients could, in some cases, have
been based on larger numbers of individuals. For in Tables A, B,
C, and D, if I throw out five sets of coefficients based on fewer than
twenty individuals, I get 89% of favorable cases. Furthermore, in
all these comparisons, the number of dimensions is quite as impor-
tant as the number of individuals ; and it is noteworthy that Table
B, with 26 dimensions and 129 individuals, gives 100% of good
cases. Finally, these comparisons are based on 335 coefficients

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BREWSTER : VARIATION AND SEXUAL SELECTION IN MAN. 49

computed from 8,551 measurements of 406 individuals belonging
to 11 different races. In Table B are four sets of coefficients, cor-
responding to four groups of individuals of the same race ; and it is
noticeable how nearly alike are these coefficients for the two sexes,
and for individuals from two different localities. This indicates
that the coefficient of variability for a particular dimension in a
particular race is a rather definite quantity. This datum is taken at
random, the choice being determined, for the most part, by the
difficulty in finding recorded measurements of both sexes. Every
coefficient computed appears in the final result, including even
those computed to find how small a number of individuals could be
taken before the method would break down so that if it should
turn out that variability is not correlated with conspicuousness, the
mistaken conclusions of this paper would be due rather to falla-
cious reasoning than to insufficient evidence or to data improperly
selected.

In discussions of variations it has commonly been taken for
granted that differences in variability are caused by the greater or
less severity of the struggle for existence. But if vai-iability is in
any way correlated with conspicuousness, this correlation can
hardly be the result of natural selection, since it is difficult to ima-
gine any connection between conspicuousness and utility. Sexual
selection will, however, account admirably for this correlation ; and
the significance of this discussion is this suggestion of the impor-
tance of sexual selection in man.

It is not clear just how sexual selection operates to increase vari-
ability instead of diminishing it as natural selection is commonly sup-
posed to do. It is possible that sexual selection is pushing men and
women in the direction of an aesthetic ideal, and in consequence
conspicuous parts of the body cannot settle down to a constant con-
dition. Perhaps variety itself is attractive, so that the individuality
which comes from looking unlike one's fellows is an advantage.

To sum up then, I have tried to show that sexual selection has
brought it about that parts of the body tend to be more variable in
proportion as they are of greater aesthetic value. There is, there-
fore, this much additional evidence in favor of Darwin's well-known
views of the importance of sexual selection in human evolution.

The data here brought together have a certain bearing on some
previous studies in variation. Prof. Karl Pearson ('97) has col-

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50 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

lected a considerable body of evidence which goes to prove that,
contrary to the commonly received opinion, men are not more
variable than women. In another paper, indeed, Lee and Pear-
son ('97) have tried to show that among civilized races men
are even less variable than women. Neither of these conclusions
b borne out by the evidence here presented. In the case of the
Pompeians — the only civilized race for which I have the data
for both sexes — the men are more variable in IB dimensions,
dhd the women in 5. Furthermore, the mean of 19 coefficients
for the men is 3.60, and for the same number of dimensions for
women is 2.68.

As for the equal variability of the two sexes in general, my
tables show 60 cases in which the men are more variable and
86 in which they are less variable than the women. Moreover,
Table 6, which gives the mean coefficient for each race of Tables
A, B, C, and D, shows that in each of the 4 cases in which the
coefficients of both sexes are given, the male is the more variable.

Lee and Pearson offer evidence ('97) to show that civilized
races are more variable than savage races. This conclusion is
contradicted by an equally large body of evidence in the present
paper. For it may be seen from Table 5 that the civilized
Pompeians are less variable than the other two savage races, the
coefficients for which are based on bone measurements; and that
of the remaining races, the savage Javanese are more variable
than any of the five civilized races except the Magyars. Thus I
venture to think that these questions of the relative variability
of men and women and of civilized and savage races are still
unsettled.

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BREWSTER: VARIATION AND SEXUAL SELECTION IN MAN. 51

LITERATURE.

Brewster, E. T. '

'97 A measure of variability and the relation of individual variations to
specific differences. Proc. Amer. acad. arts and sci., vol. 82, p. 269-280.
Carr, Lucien.
'80. Notes on the crania of New England Indians. Anniv. mem. Bost. soc.
nat. hist., 10 pp., 2 pis.
Lee, Alice, and Pearson, Karl.
'97. Mathematical contributions to the theory of evolution. On the rela-
tive variation and correlation in civilized and imcivilized races. Proc.
roy. soc. Lond., vol. 61, p. 343-867.
Marcano, G.
'93. Ethnographie prfcolombienne du Venezuela. Mfcm. soc. d'anthrop.
Paris, s^r. 2, tom. 4, p. 1-86, pi. 20.
Pearson, Karl.
'96. Mathematical contributions to the theory of evolution, 3. Regression,
heredity, and panmixia. Philos. trans, roy. soc. Lond., vol. 187, A, p.
253-318.
Pearson, Karl.
'97. Variation in man and woman in the chance of death, and other studies
in evolution. London.
Schmidt, EmiL
'84. Die antlken schftdel Pompejis. Archiv f. anthrop., bd. 16, p. 229-257,
taf. 5.
Weisbach, A.
'67. Korpermessungen « « *. Reise Fregatte Novara, Anthrop. theil,
271 pp., 8 Ub.
Weisbach, A.
'78. Korpermessimgen verschiedener menschenrassen. Zeitschr. f. ethnol.,
bd. 9, suppl.

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52 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

s
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dorso.

ventral
imensions.

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*-•

CO*

■s

in:

*» g

6 c^ i

if'

•9

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^ o

®^^ «•

ipl

•s

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face and

nose.

«s ^•

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dime

of]

and

^ s

«H O •

0'Sfc_.'d

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II-

♦o

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BREWSTER : VARIATION AND SEXUAL SELECTION IN MAN. 53

D
N

3 >
^ 8

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g

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trans-
verse di-
mensions.

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dorso-

ventral

dimensions.

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lit:

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Mean of 17

dimensions

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<N O

s s

CO CO

S S

04 CO

CO o
•o^oqi-fdi

•winT op oij

[9D

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54 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

Mean of 5
vertical
dimen-
sions.

8g§8S5SgS

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Mean of 8
transverse
dimen-
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iC^CO^CO'^-^*'^^

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ventral
dimen.
sions.

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Mean of

3 dimen- t

sions of

nose.

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Mean of
10 dimen-
sions of
face.

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Mean of
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IMft

ProoeedingA of the Boston Sooiety of Natural History.

Vol. 29, No. 3,

p. 63-75.

NOTES ON THE REPTILES A^SD AMrillBIANS OV INTERVALE,

NEW HAMPSHIRE.

By Glovkk M. Allkn.

BOSTON:
PRINTED FOR THE SOCIETY.

July, 1899.

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Ko. 3. — Notes on the Reptiles and Amphibians of Intervale^ New
Hampshire,

By Glover M. Allen, Intervale, N. H.

The observations on which this paper is based were made
almost entirely during the summer of 1898. Intervale is a small
village a few miles to the south of Mount Washington. Its fauna
is characteristically Canadian, though with a slight mixture of
Transition forms. The country in the vicinity of Intervale offers
a variety of conditions. The Saco Kiver with its broad, level
meadows or intervales is near at hand. From the edge of the
intervales the ground rises rapidly, the nearest mountains, Mts.
Bartlett and Kearsarge, being but a short distance from the river.
The woods are chiefly of white pine, beech, paper birch, yellow
birch, black spruce, and aspen. The white pines form a thin belt
between the village and the base of Mt. Bartlett. Above them
there is a well-marked belt of beeches, extending up the mountain
side. Still higher up the birches and aspens (at this elevation,
mostly the large- tooth aspen) form a mixed belt, while towards the
tops of the mountains the black spruce is the prevailing tree, with
here and there a stunted aspen (Populns tremuloides) , Near the
bases of the mountains are numerous small, clear brooks, but
larger bodies of water are few. Echo Lake, at the base of Moat
Mountain, which is just across the west side of the river valley, is
a small sheet of clear water, with a fine, sandy bottom, and so far
as I have been able to discover, has no outlet. This and Pudding
Pond, a muddy and comparatively shallow pond surrounded by a
growth of tall grass and sphagnum, are the principal ponds near
Intervale. The summers are warm but short; the winters are
cold and protracted.

During the summer of 1898, I collected nineteen species of rep-
tiles and amphibians at Intervale and vicinity, and one of the
reptiles proves to be heretofore undescribed. The list is as fol-
lows : —

1. Chrysemys picta (Herm.). Painted Tortoise.

This tortoise is found rather commonly in both Echo Lake and
Pudding Pond, and is frequently seen on bright days, sunning on

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64 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

projecting rocks or logs in the water. At such times it is shy and
will slip off and disappear in the mud, while the intruder is yet a
considerable distance away. It is amusing to watch it in its
endeavors to scramble upon a floating board. It experiences but
little difficulty in getting its fore feet upon the board, but there
the struggle begins. liaising itself to the height of its fore limbs,
with its neck stretched out forward as far as possible, it makes
frantic endeavors to get its hind feet upon the plank. After
falling back into the water several times, it finally succeeds in
gaining a hold with its hind claws, and so pushes itself forward
upon the plank. In all the specimens I have examined from
Intervale, the dorsal plates are arranged in transverse rows of
threes as in typical specimens, but it is of interest to note that
in a few specimens from eastern Massachusetts, I have seen these
scales alternating as in the western species, (7. marginata, though in
other respects these specimens were typical of C. picta. In a speci-
men taken at Intervale, late in August, 1898, the plates were in
process of peeling off. Most of them had one or two comers loose
and came off easily, when given a slight pull.

2. Thamnophis saubita (Linn.). Ribbon Snake.

On the intervale by the river, I saw what I am very positive was
a specimen of this snake, but it was so agile that it succeeded in
making its escape through the long grass.

3. Thamnophis sirtalis pallidula subsp. nov. Northern
Garter Snake.

Type locality. Intervale, New Hampshire.

Geographic distribution. From the White Mts. of N. H. and
the Adirondacks of N. Y. northward into New Brunswick and
Nova Scotia, and possibly farther.

General characters. Ground color above, olive to olive-brown ;
dorsal stripe, except at its inception, almost obsolete ; the interlinear
spots of reddish scales with narrow black edgings and black inter-
spaces. Belly, in young specimens grayish white, in adults fr^m
grayish white to light yellowish.

Description, Body rather slender, head wider than neck.
Frontal large and roughly hexagonal. Two nasals, the nostril
between. One loreal. Usually three small post-orbitals and one
large ante-orbital. Supra-labials seven ; the fifth and sixth largest.
Infra-labials ten; the fifth and sixth largest. Dorsal scales cari-
nated and arranged in nineteen rows. Anal plate entire. Gastro-

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ALLEN : REPTILES AND AMPHIBIANS OF INTERVALE, N. H. 65

ateges (from eleven specimens), 143-152. Urosteges, 117-143.
Total length, 662 mm. ; tail, 141 mm.

Color (from living specimens). Adult; ground color above,
brownish olive to olive-brown. Dorsal side of the head, olive, with
two small yellowish spots at the inner margins of the parietals.
Superior labials, light greenish white to yellowish. The dorsal
stripe, which begins at or slightly behind the posterior margins of
the parietals, is easily traceable for an inch or two as a light yellow
marking, occupying one, and two half rows of scales. From here it
may usually be traced back as far as the tail as an indistinct grayish
olive marking which, on close inspection, it is difficult to separate
with certainty from the surrounding olive-colored area. The lateral
stripes (one on each side) occupy the second and third scale rows.
In color they are from greenish yellow to ochraceous, brightest
anteriorly, and sometimes tinted with chestnut. Between the
dorsal and lateral stripes, there are, on each side, two longitudinal
series of squarish spots, best seen when the skin is distended. The
spots of the upper series alternate with those of the lower series.
Each spot involves from three to four scales of two transverse rows,
and each scale of the spot is chestnut colored, with narrow black
edgings and black interspaces. The two rows of spots are usu-
ally separated by a single longitudinal row of olive scales, which
form the sixth row in transverse series. Interspaces of other scales
above the lateral stripe are white. Below the lateral stripe, the
first row of scales and the ends of the gastrosteges are usually
light chestnut and sometimes olive-green, as in younger examples.
Beneath, from pearly white to light greenish yellow. A row of
black spots is present, on the ends of the gastrosteges. The black
of each spot may extend upward along the anterior margin of the
gastrostege and beyond so as to include the anterior margin of the
next scale or two in the rows above, but such markings do not
occur very regularly. .

Young. Similar to adult in general coloration, but the ground
color is a very pale olive, the dorsal stripe is grayish white and
rather more distinct than in the adult. The lateral stripes are very
pale yellow, sometimes hardly distinguishable from the color of the
belly, which is grayish white. A large fuscous nuchal spot on
each side.

In his original description of Coluber sirtalis, Linnaeus gives
(Syst. nat., ed. 10, 1758, p. 222) the characters as ** three greenish

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66 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

blue stripes ; narrowly striped with fuscous. Gastrosteges 150, uro-
steges 114.*' He received his specimens from Kalm and gives the
type locality as "Canada." Kalm did his collecting in northern
Vermont, along Lake Champlain, and from Montreal to Quebec.
He apparently makes no mention of this snake, in his journals
covering this region, so that it is impossible to say whether or no
he got his specimens from northern Vermont, bordering southern
Canada, or from Quebec, if indeed he got them from that region at
all, though Kalm was a careful collector and would probably have
labeled his specimens correctly. The description of Linnaeus was
doubtless taken from alcoholic specimens in which the three yellow
stripes had been turned by the alcohol to a "greenish blue." The
fact of there being three greenish blue stripes would neem to point
to the probability that he was dealing with the more southern form,
with the three bright yellow stripes, since specimens of T. s. palli-
dula^ even in life, have the dorsal stripe almost obscured, and
specimens preserved in alcohol or formalin soon lose the color of
the stripes to such an extent that in many cases the stripes would
be overlooked altogether, unless known to have been there origi-
nally. Furthermore, most of the country through which Kalm
passed, was of the Transition and not the Canadian zone, of which
paUidula is the characteristic form. The evidence, thus, as to
which of the two tj^pes, the northern or the southern, Linnaeus pos-
sessed, is about equal on both sides, though perhaps slightly in
favor of his specimens having been of the southern form. I shall,
therefore, restrict the name Thamnophis sir talis (Linn.) to the
brighter-colored form found in the Transition and Austral zones of
the East, a description of which from j^everal live specimens from
Cambridge, Mass., is as follows : — Size, proportions, and scales as
in T, s, jmliidula. Color : (young and adults) above, black, with
three greenish yellow, longitudinal stripes ; one median dorsal, two
lateral. The dorsal stripe occupies one ancj two half scale rows,
the lateral stripes occupy the second and third rows. Two longi-
tudinal series of squarish spots on each side between the dorsal and
lateral stripes, the spots of the superior row alternating with those
of the inferior. These spots are very often entirely black and can
then best be seen by pulling apart the scales, when the spots are
outlined by the black spaces between the scales composing the spot.
Occasionally one or two of the scales will have a faint reddish por-
tion in the middle, thus approaching the condition in palUdxda.

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ALLEN: REPTILES AND AMPHIBIANS OF INTERVALE, N. H. 67

The first row of scales is blackish olive to dark olive, this color
extending over the ends of the gastrosteges. A more or less per-
fect row of gastrostegal spots is present, and, as in pallklula, the
spot is frequently continuous with a black border which runs along
the anterior margin of the end of the scute, and extends up between
the scales of the first, or first and second rows. Specimens are
occasionally found which are of a browner shade than those just
described. In these there is a tendency to more or less redness in
the spots, and the spots are plainer. Some very young specimens
are lighter than described above, though generally there is but
little difference in color between old and young.

T/iamnophis sirtalis pallidula needs comparison with no other
of the sirtalis group except 2\ sirtalis proper, from which it differs
in the obscurity of the dorsal stripe, which is grayish, not yellow ;
the ground color, which is olive-brown, not black or blackish ; in
the chestnut color below the lateral stripe, where sirtalis is olive ;
in the lighter color of the belly, especially in the younger examples ;
and in the interlinear spots as previously described. The young of
pallidula are even paler than adults and are easily distinguished
from those of sirtalis proper by the gray belly and dorsal stripe,
pale olive ground color above, and the pale lateral strij)es, as well
as by the interlinear spots.

The Northern Garter snake is abundant at Intervale, where I
captured a large number. I have also taken it at Caribou, northern
Maine, and Mr. W. A. Hickman informs me that he has taken it
at Pictou, Nova Scotia. It is apparently characteristic of the
eastern Canadian zone and is usually found in or near woods. At
a short distance, the general coloring of the dorsal surface resem-
bles the color of the pine needles and dead beech leaves of its
forest home. This snake feeds largely on wood frogs {liana syU
vatica) and toads {Bufo americqnus) ^ which abound in the damp
woods. On several occasions, after having captured these snakes,
I have known them to disgorge frogs which had recently been
swallowed.

4. Natrix fasclvta sipedox (Linn.) . Water Snake.

This species I foimd only in the bog of rank grass and sphagnum
around Pudding Pond. I caught one large specimen in a " Cy-
clone" mouse trap set in a Microtias runway. The snake had
evidently been following the runway, and in attempting to pass
through the trap, had sprung it. It had died, apparently without

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68 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

a struggle, as the strong springs of the trap had come down vio-
lently just back of the skull.

5. Storeria oc^tipitomaculata (Storer). Red-bellied Snake.
Apparently this little snake was not common, though I saw

several specimens. One was seen basking in the sun on a dry
board in Pudding Pond bog, but as I approached, it quickly slipped
away among the grass. Occasionally it is found dead in the
road, having been run over by passing vehicles as it was sun-
ning itself. Dr. A. S. Packard presented me with a specimen
found, thus killed, at Intervale. Its colors were unusually bright.
The back was a blue-black instead of the usual olive-brown, and
the belly a bright red, almost scarlet.

6. LioPELTis VERXALis (DeKay). Grass Snake.

Of this species I took several specimens, and these always in
grassy places. It seems to be well distributed and rather common.
One was found dead in a grassy spot in the woods. Others were
taken at the Pudding Pond bog and on the edge of the intervales.
At Pudding Pond, I took one from under a board and brought it
home, where I placed it in a glass bottle, some three inches in
diameter. On looking at it the next morning, I found it had
nearly completed casting its skin, and only a couple of inches of
its tail still remained to be withdrawn. In the Pudding Pond
swamp I also found a cast skin, evidently of this species, in a small
bush, a couple of feet from the ground. It was twisted in and
out among the twigs, showing that it had been cast as the snake
was climbing about. I have always found this snake very gentle,
and have never had one offer to bite me. Usually, it seems to
avoid the bright light, and is often found under stones or boards.
Two, which I kept in captivity for over a month, hid, most of the
time, under a bunch of grass in their box.

7. Crotalus horrid us Linn. Banded Rattlesnake.
Although I myself have not met with this species at Intervale,

one or two are reported every summer from the vicinity. Several
have been seen or killed on Rattlesnake Ledge, a large rocky mass
on the southern slope of Mt. Bartlett. An old inhabitant tells me
that rattlesnakes were formerly common on the mountains east and
south of Intervale. One of the White Mountain guides, who has
spent most of his life among the mountains, tells me, however, that
he has never met with a rattlesnake. A specimen killed on Mt.
Bartlett a couple of years ago was examined by Dr. Packard, who

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ALLEN : REPTILES AND AMPHIBIANS OF INTERVALE, N. H. 69

telb me that it was much darker than specimens from farther south.
I have examined a couple of rattlers' skins from the Adirondacks,
and found them to be more uniformly dark than specimens from
Massachusetts, the white of the latter giving place to a dark
yellow-brown. It thus seems quite probable that the northern
rattlesnake is distinct.

8. Rana palustris Leconte. Marsh Frog.

On the broad meadows of the Saco Valley, this frog is common
along the little ponds and brooks. In £cho Lake, I found a single
young specimen in July, which still retained its tail, but otherwise
I was unable to find this species in the lake, JRana catesbiana being
the common frog there. During the summer, the Marsh Frog
may frequently be found hopping about in the grass at a consider-
able distance from the nearest body of water. On the 10th of Sep-
tember, 1897, 1 came upon a single specimen halfway up Mt. Bartlett,
and at a long distance from the nearest water. The small mountain
brooks for half or three quarters of a mile around were dried up, as
was also a small swamp near by. This swamp, of a few square
rods in extent and overgrown with bushes and sphagnum, is filled
with water during early summer, but dries up later in the season.
It is probable that this frog had been living in the marsh, and when
it dried up, was forced to seek new quarters. Other than in this
instance, I have not found the species in the woods. liana vire-
seens I did not find, though I looked for it carefully. It probably
occurs in the region.

9. Rana clamitans Latr. Green Frog.

Next to i?. aylvatica, this is probably the most abundant frog.
It occurs among the small ponds and brooks on the intervales, but
avoids the swifter waters of the river. It works its way up along
the small brooks which flow down into the river valley from the
surrounding mountains, and is not uncommonly found, along these
brooks, for some distance into the woods. Such specimens, living
in the cold clear brooks, usually average brighter in color than
those found in the open muddy ponds on the intervales. In the
summer it is not rare to find single specimens at a considerable
distance from the nearest water. I even found one hopping along
the sidewalk, one evening in August, at about one hundred and
fifty yards from the brook at the edge of the intervales. This
brook flows along the base of a steej) bank which the frog must
have climbed in the course of his wanderings. The following

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70 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

obsei-vations made on frogs of this species at Newton, Mass., are
of interest psychologically.

Several frogs living in a certain small muddy pond in a wood
were persecuted by boys throwing stones at them. This made the
frogs very shy, so that when anyone approached, they would
quickly disappear among the dead leaves at the bottom. After
having frightened them in this way, I have sometimes sat down
and waited for ten minutes or so, till they again appeared, break-
ing the surface of the water without a rij)ple, and sitting motion-
Jess as before. After they had remained thus for a few minutes,
I threw small pebbles into the water near them, but they paid no
attention whatever. P^inally I threw good-sized stones at them,
which splashed the water over them and even made them rock on
the small waves, but even then, they would often remain where they
were, apparently suspecting no danger. As soon, however, as I
arose and walked near them they saw me and retreated beneath the
surface. I have seen painted tortoises (Chryseinys jncta) show
the same indifference to stones falling near them, provided they
saw no movements to indicate danger. It would thus seem, that
these animals have learned to associate danger with the approach of
man, but after they have assured themselves that the coast is clear
they will not easily take fright unless they are warned of danger
by seeing some suspicious movement.

These frogs do not seem to feed much during the day, but at
such times sit basking in the sun. Towards evening they become
active, and may be heard every now and then splashing in the
water as they plunge after some passing insect.

10. Haxa cATEsniANA Shaw. Bull Frog.

Rather common in the small ponds on the intervales, whence its
loud notes may be heard well into July. In Echo Lake, this frog
is very common, and great numbers of young ones were found
there during July. These were in various stages of development,
most of them having acquired all four legs, but still retaining the
greater part of the tail. On one bright sunny day in July, I found
a great many under pieces of wood lying in the water near shore,
and saw others swimming along the edge of the lake. This frog,
with the exception of a very few 11, lyalustris^ seems to be the
only frog in the lake.

I caught one large specimen from a small muddy pond, and,
taking it home, kept it for a while in clear water in a large white

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ALLEN: REPTILES AND AMPHIBIANS OF INTERVALE, N. H. 71

jar. When captured, the frog was a dark muddy brown, but before
long it cast off the outer layer of epidermis, leaving a clear green
skin with a few dark olive blotches. This change of color on
putting it into clear water is suggestive of the cause for the
difference in color between specimens of H. clamitans living in the
clear mountain brooks and those living in the muddy ponds and
streams of the river valley.

11. Rana sylvatica Leconte. Wood Frog.

This is an abundant species in the cool, damp woods, and it
works its way well up the mountains. I caught one in a " Cyclone "
mouse trap in a dried-upbog near the top of Mt. Bartlett, at an
altitude of about 2,500 feet. This little bog was surrounded by
ledges, from which grew stunted spruces and balsams. This frog
is commonest in the beech woods and so closely resembles in color
the dead beech leaves that not infrequently, even after having seen
one jump, it is with difficulty distinguished from the background.
When frightened, it takes prodigious leaps in an erratic course,
and usually escapes into some hole or under a log. At night, while
walking in a damp spot in the woods, I found numbers of them
congregated in the path, where they had probably come to feed.
As I passed along, they jumped aside into the bushes. Rarely
have I heard them utter a sound in the summer, though occa-
sionally, when in the woods at night, I have detected their faint
rasping ** crau-au-4uk."

12. BuFO AMERICANU8 Lccontc. Commou Toad.

This is an abundant species from the river well up the mountains.
During the summer, very many small ones, ranging in size from one
half an inch to an inch and a half, are to be found in the woods.
The frantic struggles of these little ones as they scramble over the
leaves and twigs are very comical. Frequently they make no
progress at all, and struggle blindly to get up an almost perpen-
dicular slope, with a dogged persistence, which, however, some-
times accomplishes its purpose. Toads are abundant on the grassy
intervales, and after dark many appear in the roads made for hay-
ing carts. Apparently they are in search of insects, and are also
attracted by the warmth of the dusty roads which have all day
been exposed to the sun.

After the breeding season, the toad's song changes from a shrill,
prolonged pipe to a shorter, lower-toned note, that, at night, has a
peculiar weirdness, and almost reaches a wail. This note is heard

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mostly at evening and during the night, though I have occasionally
heard it early in the morning and in the late afternoon.

13. Hyla picKEKiNGn (Storer). Pickering's Hyla.

This delicate little frog is very abundant in the woods and groves,
especially in the damp, deciduous woods of beech, birch, and aspen,
along the bases of the mountains. It seems to live mostly well up
in the trees, whence its shrill cries are often heard. After July, it
seems to get more noisy in the woods, and may be heard at almost
all hours of the day. These frogs appear inactive during the night,
though I have heard their notes in early September until about
7.30 p. M.

They are hard to find, as they will frequently, after uttering a
few notes, stop piping and remain silent for long intervals. They
are occasionally found hopping about in the woods, and at such
times can be captured with little difficulty, though they are almost
as nimble as wood frogs. Apparently they can change their color
only to a limited extent, by making the X-mark on the back and
the other darker markings change from a rich yellow-brown to a
gray so pale as to be almost indistinguishable from the ground
color of the back. When jumping from one leaf to another, or
to any vertical surface, they will always come to rest with the
head pointing up. When placed in a bottle, and then turned so
as to be upside down, they will always turn around until the ver-
tical position, with the head up, is reached.

14. Hyla versicolor Leconte. Tree Toad.

Abundant in groves, orchards, and shade trees from the river
valley into the woods, though apparently commonest in the more
open areas between the river and the woods. During June and
early July, it is heard tooting from the trees on every side, dur-
ing the warm evenings. As the summer advances it is less often
heard, and during August and September only an occasional
note is sounded. It sings mostly during the evening and night,
though after a shower during the day, its notes may be heard. I
have also occasionally heard it in the woods at noon, on hot sunny
days.

I once found one on the intervale, squatting on a large, dark
colored rock, under an oak. It looked very much like a thick
piece of lichen and had turned a grayish white color. It was cling-
ing to the rock with head lowered, and feet tucked in close to the
body, so that at first I was completely deceived, and only on
attempting to remove the lichen, did I discover it.

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15. DiEMYCTYLUs viRiDESCEXs Raf. Common Newt.
Apparently this species was uncommon. From Echo Lake, I

took a single specimen in the larval condition, about the last of
July. Another, an adult, was found dead in a small pool on the
intervale, but other than these I found no specimens of this form.
Of the land form, " mmiatiiSy* I found, late in September, a single
specimen, lazily swimming in shallow water near the shore of Echo
Lake. It was so listless in its movements that I easily captured
it in my hands.

16. Desmogxathus FuscA . Raf. Dusky Salamander.

The commonest salamander. I found it in numbers under
old logs, stones, or leaves near the small mountain brooks, but in
no case were adults found in the streams. What I took to be the
larvae of this species, though I did not rear them through, were
common in all the pebbly mountain brooks, especially in places
where there was a bottom of granite sand and pebbles. Not
infrequently I found the adults at a considerable distance from any
brook, but in such cases they were always under old logs in cool,
damp places. When uncovered they could easily be captured, as
they seemed dazed by the sudden light. They seem to be most
active at night. I once caught three and put them in a small
bottle with a perforated cork in its mouth. During the day, they
remained contentedly in the bottle, occasionally moving about or
trying to assume an erect position on the side of the bottle. Dur-
ing the night, however, the two smaller ones managed to crawl
through the hole in the cork, and were found next morning cov-
ered with dust, on the floor. The third could not have gone
through the hole by reason of its larger size, otherwise it would
doubtless have followed the others.

None of the Intervale specimens had any perceptible membran-
ous expansion on the tail, and were, for the time, terrestrial. I
therefore carefully compared them with Copers original description
of Desmognathus ochrophaea (Proc. acad. nat. sci. Phil., 1859,
p. 124), which is as follows: — "Color above, varying from
bright, to dirty and fuscous straw-color, most specimens with an
indefinite medial row of irregular brown spots ; a deep brown line
passing through the eye and along the dorsolateral region of the
body to the end of the tail; distinctly defined along its upper edge,
fading into fuscous marblings on the sides. Belly pure white.
Susquehanna Co., Penn. A terrestrial species."

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74 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

In Bulletin 34, U. S. nat. mus., 1889, he gives a more extended
description, and mentions the light line from eye to rictus, and gives
the costal grooves as thirteen, not counting the one immediately in
front of the groin. The mandibular teeth of the males, he says, are
longer than in /usca. Through the kmdness of Mr. S. Garman, I
have been enabled to examine the specimens of Desmognathus in
the collection of the Museum of comparative zoology, which,
together with the series in my collection, makes a total of about 90
specimens from various points from Rangeley, Maine, to Georgia.
The Intervale specimens agree well with Cope's description in most
cases, but the entire series shows that there is a considerable
variation in the amount and extent of dark mottling underneath and
on the sides, and that there is no constant color difference between
/usca and ochrophaea. In the small series from Rangeley, Maine,
one specimen, a very old individual, is almost black all over, but has
the belly somewhat mottled with white, and is provided with a
membranous caudal expansion. In the same lot are other specimens
agreeing • perfectly with the description of ochrophaea. The
specimens from North Carolina are dark, with the membrane, and
a specimen from Georgia is light on the belly, but its tail has been
lost. A series of breeding specimens from Plainfield, N. J., are
colored as in ochrophaea and have the tail expansion. The series
from Intervale shows variation from light bellies to heavily mottled,
but all lack the caudal expansion. In short, an examination of the
entire series shows that the color variation is entirely individual and
seems to have no geographic bearing ; that the color differences as
well as the character of length of teeth have no weight ; and that
the only difference between the two species is the presence or
absence of a caudal expansion. This character can hardly be looked
upon as having much weight. Specimens which can easily get out
upon land after the breeding season, would doubtless, if they
remained away from the brook, suffer a reduction of the membrane,
just as the webs in certain of the Anura are reduced after the
breeding season is over. Others, which from necessity or choice
remain in the water, would retain the membrane in consequence.
Specimens of ochrophaea taken in the breeding season would
doubtless show the membranous expansion, and, indeed, the series
from Plainfield, N. J., which has the light bellies and coloring as
Cope describes, shows also the fin-like membrane on the tail. I can
not find that there is a single good character for ochrophaea.

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ALLEN: REPTILES AJSD AMPHIBIANS OF INTERVALE, N. H. 75

17. Spelerpes BILINEATU8 (Green). Two-lined Salamander.
This slender little salamander is rather common under pieces of

wood by the wet banks of little forest brooks, in mach the same
localities that Desmognathus fusca inhabits. It is agile, and when
once aroused, scrambles about with great rapidity. I found one in
the crannies of a big log, lying across a small brook. On trying to
capture it, it scrambled off the log, and, plunging into the water,
swam quickly to the bottom and hid beneath a leaf. Like other
salamanders, it will walk off the edge of a high box or table with the
utmost unconcern, and strike the floor without suffering any appar-
ent inconvenience.

18. Plethodon erythronotus (Green). Red-backed Sala-
mander.

This is a common species under old logs in the damp beech
woods, where there are one or two under almost every old log.
These logs are, in most cases, sunken slightly into the ground, so
that there would seem to be no means of egress at the sides, but
there is usually a small hole or two leading from underneath the
log down into the ground, and into these holes I have seen the sala-
manders go when pursued. Whether they make these holes or not,
I do not know, but they evidently use them to get in and out under
the logs.

The series taken at Intervale shows a considerable range in color
variation. Some specimens are coal black on the sides, with a
bright red stripe, and others are only lightly mottled on the sides
with white and dusky, the dorsal being rather dull. In this species
and the two preceding, it will generally be found that the part of
the dorsal stripe just at the base of the tail is the brightest and
most free from darker spots, and remains distinct longest when the
animals darken with age, as in the case of Desmognathns fusca,

19. Amblystoma punctatum (Linn.). Yellow-spotted Sala-
mander.

After a rain^ I obtained a single specimen of this species from
under an old decaying log in the beech woods. Further search
failed to reveal others.

Note, Since writing the above a fine specimen of Thamnophia
aaurita has been taken on the intervales.

FrinUd, July, 1899,

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FtoceddlJigs of the fioBloii SQolety of H«tur«l Mi^Lory,

Vol, "^K No, 4,

fi. 7 7 -fill.

SITMHK^ [S DiriKliA rTrLOIUlAiniA. L niE riPUNLnTLlDAB
|>K IIIK UNITKIl STATES.

llr ffAKiiv nr, N Ittitioii

BOSTON:
PHINTFhD for the 80CIKTV

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No. 4. — Studies in Diptera Cyclorhapha, 1. The Pipunculidae
of the United States,

By Garry de N. Hough, New Bedford, Mass.

This is a small family of mostly small flies, from three to five
millimeters in length. A better idea of their appearance can be
obtained from the figures on p. 473 of Professor Comstock's Man-
ual for the study of insects than by any description that can be
written.

The family contains but four genera which may be thus dis-
tinguished : —

1. Occiput scooped out, closely applied to the convex cephalic

surface of the thorax. Body, long, hairy ... 2

Occiput not scooped out, not so applied to the thorax. Body

naked or very slightly hairy 3

2. Discal cell present .... Prothecus Rond.
Discal cell absent .... Cualarus Walk.

3. Abdomen elongate, thorax with well-developed bristles

Nephrocerus Zett.
Abdomen not particularly elongate, thorax without well-
developed bristles .... Pipunculus Latr.

For an elaborate characterization of these genera see Becker's
monograph of the Eui:opean species, Berl. ent. zeits., 1897, vol. 42,
p. 25-100.

Of the four genera of this family all except Nephrocerus are now
known to occur in this country.

Chalarus. In my collection are two specimens, apparently of
different species. One was collected here by myself, the other in
Colorado by C. F. Baker. Neither is in sufficiently good condition
to use as the basis of a description.

Prothecus. Pipunculus lateralis Walk. (Dipt. Saund., p. 216)
is referred to Prothecus by Mr. Coquillett (Proc. acad. nat. sci.
Phil., 1895, p. 331). Pipunculus opacus Will. (Trans. Amer. ent.
soc, 1886, vol. 13, p. 295) also belongs here. The femora of
P. lateralis are " serrated beneath for half their length with very
small black teeth " ; the femora of P. opacus are not so serrated.

Pipunculus. By the kindness of Mr. Samuel Henshaw I have

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78 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

recently been permitted to examine the types of Loew's species and
thus to make sure of the correctness of the determinations of the
species in my collection. As far as I am aware, none of the species
described and tabulated by Professor Williston in Biologia Cen-
trali- Americana, vol. 3, have yet been found in the United States.
It is not unlikely that some of them will be found in our most south-
em states. I strongly suspect that P, reipublicae Walk, is the
same as P, fuscita Lw. and that P. tranalatua Walk, is P. subvi-
reacens Lw. Professor Williston's description of P, aridus (Dipt,
of Death Valley expedition, p. 255-250) applies exactly to P.
aubvirescens Lw., with which it is therefore probably synonymous.
I shall arrange the species in accordance with the tables in Becker's
monograph.

Division 1. Stigma wholly or partly colored; abdomen wholly
opaque ; third antennal joint usually long acuminate.

Division 2. Stigma wholly or partly colored; abdomen wholly
or partly shining ; third antennal joint usually obtuse.

Division 3. Stigma not colored at all.
Division 1.

1. Fourth longitudinal vein without an appendix,

2. Legs not wholly black, at least the knees yellow.

3. Abdomen naked, at most with a few scattered, erect, fine hairs.
5. Hypopygium (sixth abdominal segment) of varying size but

not larger than two abdominal segments together.

7. Stigma (subcostal cell) not colored its whole length.

8. Males 9

Females 10

9. Fourth longitudinal vein with a very distinct angle at its
junction with the hind cross vein.

a. Hypopygium without a cleft.

Third antennal joint obtuse . . fasciatus Lw.

Third antennal joint prolonged into a long white

process ..... subopacus Lw.

b. Hypopygium with a cleft.

Hypopygium small, shiny, only thinly poUinose; cleft
to the right of the median line . 7iigripes Lw.

Hypopygium large, opaque, thickly gray pollinose

except a spot near the cephalic border ; cleft to left

of median line . . . atlafHicvs sp. nov.

P. fasciatus Lw. In the Loew collection there is a single male.

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HOUGH: PrPUNCULIDAE OF THE UNITED STATES. 79

Opaque, black ; halteres and antennae black ; third antennal joint
short and obtuse. Dorsum of thorax opaque with brownish gray
pollen. Base of each abdominal segment black and opaque; the
remainder of the first segment densely white pollinose; remainder
of the other segments with grayish pollen which becomes whiter,
thinner, and less opaque toward the sides. Sixth segment of mod-
erate size, black. Legs black ; apices of femora and bases of tibiae
yellow or reddish yellow. Wings gray, toward base more purely
hyaline; stigma brown; veins black; small cross vein near the
junction of the basal and middle thirds of the discal cell; third
segment of costa shorter than the fourth.

P. subo2)acu8 Lw. In the Loew collection a single female.
Brownish black. Halteres yellow. Antennae black ; third joint
prolonged into a long white process. Dorsum of thorax lightly
sprinkled with brownish gray pollen. Abdomen brown black
almost shining; first segment except the cephalic margin, other
segments toward the sides and caudolateral angles whitish polli-
nose. Sixth segment of male small and without cleft. Femora
black, thickly gray pollinose, except mesal surface of posterior
femora, which is shining black. Apex of femur sometimes yellow-
ish. Tibiae vary from yellowish with a black ring to black (some-
what gray pollinose) with yellowish base. The tarsi are black or
brown with more or less yellow on the first one or two joints.
Wings grayish hyaline; stigma brown; veins black; small cross
vein at about the junction of the basal and middle thirds of the
discal cell ; third segment of costa about equal to the fourth.

P. nigripeB Lw. In the Loew collection there is a single male
without a head. Brownish black. Antennae black; third joint
short acuminate. Dorsum of thorax and scutellum brown polli-
nose, subopaque. Abdomen wholly opaque. First segment, except
at base, white pollinose. Second, third, and fourth segments vel-
vety black, each with a caudal, transverse, dark gray pollinose
fascia. Fifth segment brown pollinose, subshining, whitish polli-
nose on the sides. Sixth segment black, rather shiny, thinly gray
pollinose; its cleft considerably to the right of the median line.
Legs black. Femora grayish pollinose to a varying extent except
the mesal surface of the posterior femora, which is shining black ;
the extreme tips of the femora are yellow. Flexor surface of
anterior and middle femora with rows of tiny black spines. Tibiae
yellow at base to a varying extent. First one or two joints of the

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80 PROCEEDINGS : BOSTON SOCIETY NATUHAL HISTORY.

tarsi yellow or brownish yellow to a varying extent. Wings gray-
ish ; stigma brown ; veins black ; small cross vein at a point
apicad the junction of the basal and middle thirds of the discal cell,
but before the middle of that cell ; third costal segment about twice
as long as the fourth. Ilalteres black except base of peduncles
yellowish.

P, atlanticua sp. no v. Two males and three females. New
Bedford, Mass., Hough; Montgomery Co., Pa., C. W. Johnson.
Length 3 mm., length of wing 5 mm. Brown black, mostly gray
pollinose, with wholly yellow legs and very distinctly banded abdo-
men. Antennae brown with very fine whitish pubescence, third
joint short, oval, obtuse. Dorsum of thorax and scutellum black
brown pollinose, toward the sides and on the humeri more grayish.
Pleurae, metanotum, and coxae gray pollinose. Legs wholly yellow,
appearing more or less silvery gray pollinose according to the inci-
dence of the light, except the tips of the tarsi, which are black.
The four anterior femora on the flexor surfaces and all the tibiae
have rows of extremely minute black spines. Abdomen wholly
opaque. First segment wholly gray pollinose except a narrow
black cephalic border. Second segment wholly gray pollinose, the
pollen thinner on the middle of the segment, so that in some speci-
mens there seems to be a faint median, transverse, black brown fascia.
Third, fourth, and fifth segments wholly gray pollinose, except a
cephalic, transverse, blackish brown fascia which gradually fades out
toward the sides and a median, caudal, blackish brown spot which
is connected with the cephalic fascia. On the dorsum of the abdo-
men the relative widths (measured cephalo-caudad) of the blackish
brown fascia and the gray pollinose portion vary in different speci-
mens, sometimes the one and sometimes the other being the wider;
the sides of the segments are, however, wholly gray pollinose. The
sixth segment is wholly gray pollinose except a small brownish
black spot cephalad; this segment is rather large and is divided
into three very unequal portions by two dorso-ventral clefts, both of
which are to the left of the median line. The one nearest the
median line is about halfway to the lateral border, while the one
furthest laterad cuts off not over one sixth of the segment.
Remainder of genital apparatus yellow. Halteres with yellowish
brown peduncle and black knobs. Wings grayish hyaline, veins
black; small cross vein at about the junction of the basal and
middle thirds of the discal cell ; third costal segment about equal to
the fourth.

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HOUGH: PIPUNCULIDAE OF THE UNITED STATES. 81

10. Ovipositor straight.

11. Sixth abdominal segment not furrowed.

a. The lanceolate terminal piece of the ovipositor not
longer than the fifth abdominal segment

atlanticus sp. nov.

b. The awl-shaped terminal piece of the ovipositor as long
as the second, third, fourth, and fifth abdominal segments
together ...... subopacus Lw.

These females are like the males except for the sexual differences.
As far as I know, the females of P. fasciatus and P. 7i%gripes are
unknown.
Division 2.

Dorsum of thorax with delicate but distinct hairs . . 1

1. Abdomen shining black or bronze with opaque black fasciae
or spots on the cephalic borders of the segments ; males . 2
Abdomen shining black without opaque black fasciae but with
gray side spots ; mostly females 8

2. Fourth longitudinal vein not interrupted ... 3

3. Dorsum of thorax poUinose on the cephalic half without any
clearly defined boundary line between the poUinose portion
and the shining caudal portion 4

4. Abdomen with gray side spots ..... 5

5. Abdominal segments with moderately broad opaque black
fasciae at their cephalic borders.

a. Tibiae largely brownish black . . chifpilatus Lw.

b. Tibiae yellow with scarcely a suggestion of brown ; hind
tibiae rather strongly curved . . . fuscus Lw.

8. Dorsum of thorax more or less poUinose . . . 11

11. Tibiae between a brown and a yellow, seeming now one, now
the other, according to the incidence of the light. Awl-shaped
terminal piece of ovipositor straight . nitidii'entris Lw.

P./uscus and P. nitidiventris belong to the difficult group of
P. cUer Meig. It is not unlikely that they will prove to be the
two sexes of one species.

P. cingulatus Lw. In the Loew collection a single male.
** Grayish black. Antennae wholly black ; third joint short, rather
obtuse. Dorsum of thorax opaque with brownish cinereous pollen ;
pleurae sprinkled with whitish cinereous pollen. Scutellum black,
moderately shining. Metanotum white ])olIinose. First abdomi-
nal segment white pollinose with a median black spot ; the rest of

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82 PROCEEDINGS: BOSTON SOCIETY NATUBAL mSTORT.

the abdomen black, shining, with white pollinose sides, each seg-
ment with a slender, basal, dark cinereous pollinose fascia. Fe-
mora black except the yellowish apices," hind femora shining black
on mesal surface, the others not at all shining; "tibiae brown
black, basal third of all and apex of the anterior ones reddish testa-
ceous ; tarsi reddish testaceous, towards the apex brown, last joint
black or black brown. Wings slightly brownish cinereous, stigma
brown."

P, fuscuH Lw. In the Loew collection three males. Black
brown. Antennae black, third joint short, rather obtuse. Dorsum
of thorax brown pollinose, opaque, towards the caudal margin sub-
shining. Scutellum black, shining. Metanotum and first abdominal
segment, except its base which is black, cinereous pollinose ; second,
third, fourth, and fifth segments velvety black on their cephalic
halves and metallic, shining bronze-colored on their caudal halves
(a favorable incidence of light is necessary clearly to make out the
extent of the velvety black portion); the black of the second seg-
ment is to a certain extent cinereous pollinose, and each segment
has on the sides an indistinct grayish pollinose spot; the fifth
segment is much longer than the preceding ones ; the sixth is much
larger still, not symmetrical, and wholly shining bronzy except for a
widely interrupted transverse fascia on its cephalic border. Tibiae
yellow with scarcely a suggestion of brown ; hind tibiae rather
strongly cur\'ed. Femora black except base and apex which are
yellow ; lightly white pollinose except the mesal surface of the hind
ones, which is shining black ; all of them with a certain amount of
white pile, the pile on the posterior surface of the middle femora
much longer and denser than elsewhere ; flexor surfaces of all with
rows of very minute black spines. Wings cinereous; stigma
brown ; small cross vein at junction of basal and middle thirds of
discal cell; veins black; third costal segment longer than the
fourth. The relative lengths of the third and fourth costal seg-
ments in this and the following species are very difficult to deter-
mine on account of the extremely acute angle at which the first
vein meets the costa, the brown color of the stigma, and the rather
large size of the veins, so that the exact point of junction of the
first vein and costa is hard to see.

P. nitidwentrls Lw. In the Loew collection a single female.
Blackish cinereous. Antennae black brown, thii-d joint with the
apex short acuminate. Dorsum of thorax opaque with whitish cine-

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HOUGH: PIPUNCULIDAB OF THE UNITED STATES. 83

reous pollen, in the middle brownish cinereous. Scntellum black,
moderately shining. Metanotum whitish pollinose. Abdomen
wholly black and shining; first segment except at base white
pilose ; second very slightly sprinkled with cinereous pollen so that
it is less shiny than the following segments, its sides and caudal
margin almost wholly white pollinose ; third segment with a cine-
reous pollinose stripe, dilated cephalad, obsolete caudad ; the sides,
the caudo-lateral angles, and the lateral portions of the caudal
margins of segments 3, 4, and 5 are white pollinose ; the sixth seg-
ment has its lateral margins and its caudo-lateral angles white pol-
linose. The tibiae are between a brown and a yellow color, seem-
ing now one and now the other according to the incidence of the
light. Femora black except the extreme bases and apices, which
are yellow ; they are wholly pollinose except the mesal surface of
the posterior pair, which is shining ; I can see no spines or pile on
either femur. The awl-shaped terminal piece of the ovipositor is
straight. Wings hyaline ; veins black ; stigma brown ; small cross
vein at junction of basal and middle thirds of discal cell; third
costal segment shorter than fourth.
Division 3.

1. Stigma not colored 2

2. Small cross vein at or apicad the middle of discal cell . 3

Small cross vein at basal fourth or tifth of discal cell . 10

Small cross vein before the middle but beyond the first third

of discal cell 22

3. Eyes of the male in contact near the middle of front for a
greater or less distance ....... 4

4. Abdomen, especially the fifth segment, with short, erect black
bristles 5

Abdomen with delicate whitish hairs or almost naked . 6

5. Humeri yellow. Ilypopygium asymmetrical ; looked at from
the left it is as long as the fifth sogment, from the right it is
shorter than the fifth ; on its caudal end is an oblique oval
depression which is usually very distinct. Legs black, grayish
pollinose ; apices of femora, basal third of tibiae, and tarsi
except last joint yellow; hind femora shining black on the
mesal surface. Wholly greenish black, lightly gray pollinose,
pollen thickest on first and fifth abdominal segnuMits. Anten-
nae black, very delicately whitish pubescent: third joint with
a rather long drawn out white point. Wings hyaline ; veins

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84 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

black; third costal segment one half the fourth. Length 3.5

to 4 mm similis sp. nov.

This species is very like P. sylvaticus Meig., from which its pol-

linosity and the form of its hypopygium clearly separate it. The

terminal segment of the ovipositor is straight, awl-shaped, and about

as long as the hind tibiae. I have eleven males, collected by G. R.

Pilate at Tifton, Ga., in Oct. and Nov., and two females, collected

by C. F. Baker in Alabama.

6. Tarsi yellow, only the last joint black .... 8
8. Humeri yellow; front of male very small, silvery pollinose,
without depression. Thorax and scutellum shining greenish
black, dusted with brown pollen in the middle and towards the
cephalic border. Hypopygium small, hardly half as long as
the fifth segment; its rima oblique and to the right of the
median line. Terminal piece of ovipositor straight and about as
long as the hind tibiae. Wings hyaline ; veins black ; fourth
costal segment hardly twice the third. Abdomen greenish
black, shining. Legs black ; the very apex of the femora and
base of the tibiae yellowish ; tarsi brownish yellow, toward
the tip black. Halteres with yellow knob and brownish ped-
uncle aubvtrescens Lw.

In the Loew collection a single rubbed male. I have numerous

specimens, of both sexes, from New Bedford, Mass., Tifton, Ga.,

and Opelousas, La.

10. Males 11

11. Eyes not in contact, front not swollen.

Shining black, lightly white pollinose. Abdomen with a large
yellow spot on each side; fifth and sixth segments large. Legs
and coxae yellow except the basal two thirds of anterior coxae
black. Length 4 mm.

Face and ventral half of front silvery pollinose. Antennae
black, third joint yellow and very finely white pubescent, it-s apex
prolonged in a white point. Thorax shining black, very lightly
white pollinose especially near the humeri. Scutellum shining
black. Metanotum and ])leurae somewhat whitish gray pollinose.
First abdominal segment black with a transverse yellow fascia at
the cephalic border ; second segment black except the caudo-
lateral angles, which are yellow; incisure between second and
third segments yellow ; third and fourth segments black on the
middle of the dorsum, their sides almost wholly yellow ; incisures

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HOUGH: PIPUNCULIDAE OF THE UNITED STATES. 85

between third and fourth and between fourth and fifth segments
yellow ; fifth segment wholly shining black, much longer and wider
than fourth segment, beset with a few delicate, scattered, white
hairs; sixth segment wholly black, somewhat grayish pollinose,
quite as large as the fifth segment, not symmetrical, being larger on
the left side, no visible cleft or depression on its apex. Wing
grayish hyaline ; veins black, yellow towards the base ; small cross
vein at basal fourth or fifth of discal cell ; fourth costal segment
much longer than the third. One male ; Horse Neck Beach, Mass.,
Aug. 6 flavomacidata sp. nov.

12. Brown black, poUinose. Abdomen with segments one and
five densely gray pollinose. Femora black, gray poUinose. Tibiae
brownish yellow. Tarsi yellow except last joint black. Length
3.5 mm.

Eyes in contact. Face and the small, smooth frontal triangle
silvery white. Antennae black, third joint silvery with a rather
short sharp point. Dorsum of thorax and scutellum brown polli-
nose tending to gray pollinose at the sides and near the scutellum.
Humeri yellow. Pleurae gray pollinose. Metanotum densely gray
pollinose. First abdominal segment black on cephalic half, densely
gray pollinose on caudal half; segments two, three, and four are
blackish brown, very lightly gray pollinose on the dorsum, the
pollen becoming gradually much thicker towards and on the sides ;
fifth segment very densely gray pollinose ; on it there is visible in
some lights a suggestion of a median, cephalo-c^udal brown stripe ;
sixth segment shining black, only very slightly pollinose save at
sides and on venter, about as large as the fifth segment, very
slightly asymmetrical ; no rima or depression can be seen ; segments
two, three, and four are of equal length, fiye is distinctly longer.
Femora black, their extreme apices yellow, wholly gray pollinose
except the mesal surface of the posterior, which is shining black.
Tibiae appear to vary somewhat in color, according to the strength
and incidence of the light, between a yellow and a brown; the
anterior tibiae usually look brown except the extreme base which
is yellow; the other tibiae look yellowish brown. Tarsi: anterior
brown with yellow base ; the others yellow with black tips. Wings
grayish hyaline; veins black; small cross vein a little apicad the
junction of the basal and middle thirds of the discal cell; third
costal segment one half as long as the fourth. Halteres blackish
brown, the middle of the peduncles yellowish cUbo/asciata sp. nov.

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86 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

One male, Opelousas, La. Collected by Mi-. G. R. Pilate in
May, 1897.

In the last two species, as in P.fuacuSy the relative lengths of
the third and fourth costal segments are very difficult to determine ;
moreover I cannot be quite sure that there is not a very little brown
color in the stigma at its extreme apical angle. There is, however,
so little of this color, if any, that there can be no impropriety in
placing the species in this third division.

Only about ninety species of this family are known, of which fifty-
eight are European. Undoubtedly there are many more species to
be discovered.

Printed, July, 1800,

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Ptoca^dine* of tile Bostou BoQietf of IVaturai Hislorj.
Vol, 2% No, 5,

riiXTJnrU'1'IONS KKHM THK <»UAV IIKRBARII'M UF JlARVArd)

Rt B L Rotii\sijs imo *[. M. (iiff;f,Mwt%

BOHTON:

PRINTED FOR THE SOCIETT.

AutirsT, 1890.

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No. 5. — Contributions from the Gray Jlerbarium of Harvard
University. Nein Series. — No. 17.

By B. L. Robinson and J. M. Greexman, Camiiridge, Mass.

1. REVISION OF THE GENUS GYMNOLOMIA.

The genus Gymnolomia, so far as known, is exclusively Ameri-
can and extends from the northwestern parts of the Unit€»d States
to Brazil. More than half its species, however, are confined to
Mexico and Central America. The genus in its present circum-
scription has never been monograj)hed. Kunth when founding it
in 1S20 described 4 species. De Candolle in the Prodromus, 5, 5(51
(1^86) changes rather arbitrarily Gymnolomia to Gymnopsis under
which 10 species are described of which several have since
been referred to other genera. In 1^78, Bentham and Hooker f.
ascribe to the genus Gymnolomia 16 species. Baillon in his His-
toire des plantes regards the genus Gymnolomia as of doubtful
value. Hoffmann in Engler and Prantl's Nat. Pflanzenfamilien
maintains the genus and ascribes to it 20 species. Aside from the
brief citations of the Index Kewensis by far the best bibliographic
enumeration of the Gymnolomias is that of Hemsley (Biol. Cent.-
Amer. Bot., 2, 161) who cites literature and stations for 15 species,
found in Mexico and Central America.

The present revision, including 87 recognized species and several
varieties, has been chiefly based upon the specimens in the Gray
herbarium, now largely supplemented by the incorporation of the
Klatt herbarium. But through the kindness of Mr. Coville and
Dr. Rose, the representation of the genus in the U. S. national
museum has also been examined with profit.

The revision of the genus has presented three chief difficulties,
viz.: — 1. The distinction of our own western and southwestern
forms, which have, notwithstanding great diversity of foliage, pu-
bescence, size of heads, and duration, been loosely grouped under
G, niidtiflora. 2. The exact interpretation of the Central Ameri-
can G. suhflexuosa^ Benth., upon which must depend the varietal
and specific nomenclature of some of the nearly related 8])ecies.
3. The explanation and proper treatment of several large-headed

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88 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY

Mexican species which although they differ conspicuously among
themselves, closely simulate a parallel series of Viguieras. As
examples of this parallelism may be mentioned

G. ensifolia with the habit of V. blepharolepU.

G. meyacephala^ var. sitnulajis with the habit of V. exctlsa.

G. decurnbe7i8 with the habit of V, (jhiesbrtfjhtii.

G, ghiesbreghtii with the habit of V. buddleiaefomiis.

This simulation has various degrees of closeness. In G. yfiies-
breghtii and V. buddlti(teforniis^ although the similarity is striking,
the heads are always perceptibly smaller in the Viguiera. G. me-
gacephala var. simulans and G. deciunbefitiy however, so closely
resemble V. excelsa and F! yhitHhreghtii res[)ectively, that no
satisfactory external characters have as yet been found by which
to separate them. The examination of the achenes, however, at
once reveals striking differences. The achenes are in the Gymno-
lomias more compressed, quite glabrous, and completely destitute
of pappus. In the Viguieras on the contrary they are thickish,
obtusely 4-angled, upwar<lly villous, and provided with two stout
persistent awns and several short intermediate scales. Were the
pappus difference the only one, these viguieroid Gymnolomias
might logically be reduced to ^\format epitpposne " of their pappus-
bearing counterparts in Viguiera, there being in Galea and else-
where undoubted examples of the presence and absence of pappus
in what must be regarded as conspecific types. In Gymnolomia,
however, the difficulty is increased by the presence of other although
slight differences in the form and pubescence of the achene.

The writers have carefully considered the possible readjustment
of generic lines in such a manner as to bring together the species
of similar habit, but they have failed to find any way in which this
could be accomplished without the union of heterogeneous ele-
ments or an inordinate multiplication of genera founded upon
rather trivial characters. It has accordingly seemed best to main-
tain the genus Gymnolomia in its traditional interpretation,
although there may be some artificiality in the classification of
the large-headed forms. Further collections will doubtless throw
much light, if not upon the genetic affinities, at least upon the
validity or inconstancy of the characters by which they are now
separated.

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ROBINSON & GREENMAN : CONTRIB. GRAY HERBARIUM. 89

GYMNOLOMIA, HBK. (Name from yv/ivo«, naked, and
XCifioj margin ; taken by some to refer to the rays destitute of pis-
tils, by others the achenes lacking pappus.) Heads small, medium-
sized, or large, heterogamous, many-flowered, pedunculate, solitary
or more often 3 to many at the ends of the branches. Involucre
hemispherical or subcylindric ; the bracts mostly numerous, in 2 to
4 series, very variable in size and thickness, the outer mostly nar.
row; receptacle elevated, conical, chaffy; pales conduplicate,
entire or somewhat 3-toothed. Ray-flowers (5 to 20) in a single
series, neutral ; ligules linear to elliptic-oblong, exserted and spread-
ing, entire and slightly toothed at the tip, yellow or very rarely
crimson ; disk-flowers numerous, perfect, regular, concolorous ; tube
short, cylindrical or somewhat dilated at the base, usually puberu-
lent ; throat cylindrical or narrowly campanulate ; the limb 5-
toothed. Achenes of the ray- flowers abortive and empty, of the
disk-flowers obovoid, thickish, but more or less compressed laterally
or 4-angled, rounded at the summit ; pappus none or rarely present
as a short cup or ring of 2 to 4 laciniate scales. Nov. gen. et
spec, 4, 217, t. 373, 374; Gray, Proc. Amer. acad., 5, 182; Syn.
fl., 1, pt. 2, 66, 269; Benth. and Hook, f.. Gen., 2, 363; HerasL,
Hiol. Cent.-Amer. Bot., 2, 161; Baill., Hist, pi, 8, 211 ; Hoffmann
in Engl, and Prantl, Nat. Pflanzenf., 4, ab. 5, 233. Gymnopsis,
DC, Prodr., 5, 561, in part. Heliomeris, Nutt., Journ. acad. Phila.,
ser. 2, 1, 171. Zaluzania, Sch. Bip., Flora, 1861, p. 553; 1864,
p. 216, in part. — Nearly 40 species, chiefly perennial herbs, in
habit passing almost imperceptibly from plants of the Perymenium
type to others of the Tithonia type. The following arrangement is
believed to show approximately the natural aflinities of the species.

Subg. 1. CALANTirARiA. Corolla-tube in the disk-flowers con-
siderably dilated at the very base, the expansion, thus developed,
forming a sort of cap over the summit of the achene : stems shrubby,
except perhaps in G. tripartita : heads medium-sized ; scales of the
involucre mostly narrow.

• Leaves rather small, opposite, entire or merely undulate : scales of the
involucre oblong, obtuse.

1. G. GREGGii, Gray. Closely branched shrub : leaves opposite,
ovate, obtuse, cuneate to short petioles, pale green above, white-
tomentulose beneath, 2 to 3 cm. long (incl. petiole), 1 to 1.8 cm.
broad; peduncles long, mostly solitary at the ends of the branches.

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90 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

Proc. Amer. acad., 15, 36. — Northern Mexico, Gregg^ no. 382;

and on limestone hills Cameros Pass, Coahuila, Pringle, no. 2387.

♦ * Leaves alternate, larger, subentire.

2. G. ciXERASCENs, Benth. and Hook. f.. Dichotoraous under-
shrub with brownish purple soft-pubescent leafy stems terminating
in compound irregular many headed corymbs : leaves ovate-elliptic,
5 to 6 cm. long, 3 to 4 cm. broad; the upper oblong, attenuate at
each end, acute, minutely serrulate, subtriplinerved, pubescent
above : heads borne on white clavate peduncles : ligules (white ?)
neutral : achenes glabrous, without pappus. — Gen., 2, 362 ;
Hemsl., 1. c, 161. Zcduzania cinerascens, Sch. Bip., Flora, 1864,
p. 219, whence above desc. is compiled. — S. Mexico, Mineral del
Monte, Ehrenherg^ no. 346. We have not been able to see speci-
mens of this species but from Schultz statement that the corolla is
cucullate over the achene there can be little doubt that it is to be
referred to Subg. Calanticaria.

* ♦ * Leaves lobed, scales of the involucre lance-linear, acute to attenuate.

3. G. piNNATiLOBATA, Bcnth. and Hook. f. Shrub: branches
covered with a fine white at length deciduous pubescence : leaves
pinnately several-lobed, pale green above, white-tomentulose
beneath, 3 to 6 cm. long (including the winged petiole) , half as
broad ; lateral lobes blunt, the lower large, the upper smaller. —
Gen., 2, 364; Hemsl., 1. c, 163. Zaluzania pinnatilobata^ Sch.
Bip., Flora, 1864, p. 219. — South Mexico, Tehuacan, Liehmann,
no. 384, Pringle, no. 6252; Cordillera of Oaxaca, Galeotti, no.
2124, ace. to Hemsl. ; limestone mesas, San Antonio, Oaxaca, Prin^
gle, no. 5731. Mr. E. W. Nelson's no. 1638 from Oaxaca forms a
puzzling intermediate between this and the otherwise well-marked

G, tripartita..

4. G. TRIPARTITA, Robiusou and Greenman. Stems smooth and
glabrous : leaves deeply and palmately 3-parted ; segments narrow,
oblong, acute, often bluntly lobed near the base, the lateral some-
times very short or in upper leaves wanting, paler but green be-
neath: heads numerous, slender-peduncled in an open corymb. —
Amer. journ. sci., ser. 3, 50, 154. — Oaxaca, Cuicatlan, X. C. /Smith,
no. 239 ; Jayacatlan, X. C. Smith, no. 386.

5. G. TEXiriFOLiA, Benth. and Hook. f. Low shrub, 1 m. or
more high : leaves numerous, deeply 3-fid or pinnatiiid with lobes
few, linear to lance-linear, acute, green above, finely white pubes-

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ROBINSON & GREENMAN : CONTRIB. GRAY HERBARIUM. 91

cent beneath, the margins re volute : peduncles long, solitary at the
ends of the branches; bracts of the involucre attenuate. — Gen., 2,
364; Hemsl, I.e., 168; Gray, Syn. fl., 1, pt. 2, 269. Ildiomeris
tenuifolia, Gray, PI. Fendl, 84; PI. Wright, 1, 107; 2, 87.—
Southwestern Texas, rocky cliffs of Turkey Creek and beyond the
Pecos, Wright^ no. 329; rocky hills near Eagle Springs, Wright,
no. 1223, Sutton Hayes^ no. 444 ; at Presidio, Ilavard^ El Paso,
6^ay, Jones, no. 4347; Santa Maria, NeaUey, no. 163; and Che-
nantes Region, Nealley^ no. 576 ; New Mexico in the Organ Mts.,
Wooton, no. 442; Mexico, Tamaulipas, Berhmdier^ nos. 814, 2234,
E. W, Nelson, no. 4507; Saltillo, Gregg^ nos. 21, 125; Coahuila,
Palmer, nos. 620 to 624 (coll. of 1880), Pringle, no. 148; Chihua-
hua, Thurher, no. 834 ; San Luis Potosi, E, W. Nelson, no. 4532.

Subg. 2. EuoYMNOLOMiA. Corolla-tubc in the disk-flowers
cylindrical or slightly enlarged at the base ; leaves (often serrate)
not lobed ; stems mostly herbaceous.

* Heacla usually numerous, pedunculate, small or more often medium-sized
(disk 7 to 12 mm. broad excl. of the rays); involucral scales linear, acutish
to attenuate : leaves mostly lance-linear to linear, rarely oblong.

-♦- Outer involucral bracts herbaceous, attenuate (1.1 to 1.7 cm. long) coarsely
ciliate on the margin, otherwise nearly or quite glabrous : ligules 5 to 9,
elliptic, showy : Georgia.

6. G. PORTEBi, Gray. Tall slender erect corymbosely branched
annual with lance-linear to linear chiefly alternate leaves coarsely
ciliate near the base; those of the stem 6 to 12 cm. long: disk nar-
rowly conical ; pales striate, entire, pungent. — Proc. Amer. acad.,
12, 59; Syn. fl., 1, pt. 2, 269; Meehan, Native fl., ser. 1, 2, 137, pt.
35 ; Chapm., Fl., ed. 3, 251. Biidbeckia ? porteri. Gray, PI. Fendl,
83; Chapm., FL, ed. 3, 228. — Growing on scanty humus in hollows
and crevices of granite rock on the summit and slopes of Stone
Mountain, Georgia, where curiously local, Porter (type, in herb.
Gr^^y) , Bavenel, Ilendee, Otnby, J, 2>. Smith, Cnrtiss, no. 1434,
Small ; fl. August to October.

t- t- Outer involucral bracts (rarely elongated) usually appressed-pubescent
or even hoary : ligides 10 to 14, linear or oblong : western U. S. and Mexico.

7. G. MULTiPLORA, Bcuth. and Hook.f. Perennial : stems usually
several (1.5 to) 9 or 12 dm. high, from a thickish lignescent stock,
finely striate and covered with a very short-appressed grayish
pubescence: leaves lanceolate or linear-lanceolate and acutish to
lance-oblong or oblong and obtuse, 4 to 7 cm. long, 4 to 20 mm.

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92 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

broad, attenuate at the base, finely pubescent, becoming scabrous
especially above : beads (exclusive of rays) 1.2 cm. in diameter;
involucral scales canescent with appressed hairs. — Benth. and
Hook, f., ace. to Rothr. in Wheeler Rep., G, IGO, as to synon. ; Gray,
Syn. ft., 1, pt. 2, 269 only in part. Jleliomeria multiflora, Xutt.,
Journ. acad. Phila., ser. 2, 1, 171 ; Wats., Bot. King exp., 170, at
least in great part. — Rocky mountains and plains of Idaho, Palmer^
no. 422 ; Wyoming, Hay den, no. 34, 7\ceedy^ Btirglehaus^ A.
Nelson^ nos. 39, 1064, 2663; Colorado, Jiothrock\ no. 551, Parry ^
no. 420, Greene, no. 194, Jones^ (\ S. Sheldon, nos. 152, 468,
Cowen (low and many-stemmed), Miss EasHnood (coll. July, 1H89);
Utah, Jones ^ nos. 5S20, 599(> ; Nevada, in the Uintas, Watson,
no. 606 ; New Mexico, Wooton, no. 4S4 ; California, Ganibel, Co-
ville and Eunsto/i, no. 806; and in the Rocky Mountains, without
locality, by Gordon, Burke, and by Frhnoyit, no. 121. Toward the
southwest this species passes into somewhat narrower- leaved forms
(Utah, Ward^ nos. 647, 652, Jones^ no. 5996 r; Arizona, Palmer,
coll. of 1869 without numbqr and no. 241 coll. of 1877, Knoxrl-
ton, no. 6 ; S. E. California, Purpus, no. 5025), which, however,
retain their perennial character.

8. Q*. longifolia. Erect paniculately branched annual : stem
single, 6 to 12 dm. high, finely pubescent with appressed hairs:
leaves lance-linear or oblong-linear, narrowed at both ends, those
of the stem 8 to 11 cm. long, 5 to 10 mm. broad, entire or obscurely
and remotely crenate-serrate, finely pubescent on both surfaces,
becoming very scabrous and tuberculate-hispid above, often ciliate
toward the base ; peduncle and involucral bracts covered with fine
appressed pubescence; heads numerous, in size and floral characters
essentially like those of G, mvltiflora. — G. multiflora, Hemsl.,
1. c, 162, as to Mex. pi. ; Gray in Wats., Proc. Amer. acad., 21, 432 ;
Syn. fl., 1, pt. 2, 269, in part; not Benth. and Hook. f. Ileliomeris
multiflora, Gray, PI. Wright., 1, 107; 2, 87, in part, not Nutt. —
W. Texas, Wright^ no. 328, Jlamird, Nealley, no. 432 (depaupe-
rate), Dr. Smart, no. 422 ; New Mexico, ** on pine hills between
the copper mines and the mimbres,*' Wright, no. 1221 ; Burro Mts.,
Pushy, no. 172; Arizona, Rucker's Valley, Lemmon^ nos. 345,
2765, and 383 (a form with double flowers); Nagle's Ranch, Jones,
no. 6054 p. ; Mexico, between San Luis Potosi and Tampico, Palm-
er, no. 1102; hills and plains near the town of Chihuahua, Pringle,
no. 615; S. W. Chihuahua, Palmer, no. 392; Strawberry Valley,

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ROBINSON & GREENMAN : CONTRIB. GRAY HERBARIUM. 93

Chihuahua, Ilartman^ no. 777 ; vicinity of Durango, Pahner^ no.
754 ; Chiapas, Ghiesbreyht^ no. 68. Types in herb. Gray.

9. Q*. annua. Annual, slender, often branched from the base,
3 to 5 dm. high, copiously branched above : leaves linear or nearly
so, finely appressed-pubescent and even somewhat canescent,
slightly ciliated at the base or not at all, the cauline 4 to 7 cm.
long, 2 to 3 mm. broad, the rameal smaller : pubescence on the
peduncles and on the involucral bracts fine and apj)re8sed ; heads
rather small, usually 7 to 10 mm. in diameter (exclusive of rays):
rays 10 to 14, yellow, 4 to VI mm. long : otherwise like G. niiilti'
flora. — G. inultiflora. Gray, Syn. fl., 1, pt. 2, 2(59, in part, not

Benth. and Hook. f. G. multiflora,, var. annua^ Jones, Proc. Calif,
acad. sci., ser. 2, 5, t)9S. Jleliomeris multiflora, Gray, PI. Fendl.,
H4, as to pi. of Wislizenus ; PI. Wright., 2, S7, in part, not Nutt. —
W. Texas, Wright, no. 834, Pope ; New Mexico, on sides of moun-
tains near the copper mines, Wright ; Arizona, on sandy river bot-
toms, near Ft. Whipple, Coues and Palmer^ no. 559, at Patano,
Pringlt ; near Defiance, Mursh, no. 229 ; Arizona, Knotrlton^ no.
2S8, Wilcox, no. 455, Mothrock^ no. 779 ; Mexico, at Llanos,
Wislizenus \ in Sonora, Wright, r\o. 1220, F. E. Lloyd, nos. 410,
411. Distinguished from G. muUifloni by its annual root, much
narrower leaves, somewhat smaller heads, shorter rays and dis-
tinctly Sonoran range ; from G, hispidn^ var. ciliaUt less satisfac-
torily by the different pubescence of its leaves and involucral
scales.

10. O. hispida. Probably annual, 6 to S dm. high, densely
hispid and hoary throughout with long stiff white hairs, those of
the stem and peduncles widely spreading : leaves alternate, narrow,
linear, 5 to 7 cm. long, 3 or 4 mm. broad, thickish, channeled
above: involucral bracts considerably exceeding the disk, hirsute
and hoary: flowers as in G, multiflora. — Heliomeris niultifloruy
var. hisjnda, Gray, PI. Wright., 2, H7, as to ty})ical canescent form.
— Low damp soil, near Sta. Cruz and San Bernardino, Sonora,

Wright, no. 1220 in part. In the almost shaggy pubescence of
the leaves and copious spreading pubescence of the peduncles this
form differs rather markedly from the others here enumerated, but
it certainly passes to

var. ciliata. Slender annual, commonly branched from the
base, 3 to 6 dm. high : stems with sparse spreading pubescence or
nearly glabrous below: leaves linear, green, conspicuously ciliate

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94 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

for the greater part of their length : pubescence of the peduncles
sparing, mostly appressed ; involucral scales ciliate but scarcely
pubescent otherwise. — Ileliomeris mxdtijiora^ var. hispida^ Gray,
1. c, in part, i. e., as to greener forms. — S. Utah at Beaver, Palmer,
no. 245 ; New Mexico, Zuni Mts., Sitgreaves -£>/>., San Antonita,
Bigdow in part ; S. California at Sta. Monica, Hasse ; Sonora, Mexico,
Wright, no. 1222 in part.

* * Heads small or mediuui-sized ; involucral scales linear and acute or the
inner narrowly oblon^^ and obtusish : leaves ovate or ovate-lanceolate :
perennials so fai- as the duration is known.

■•- Heads small and numerous, corymbose at the ends of the branches : leaves
petioled (shortly so in var. abbreviata).

■^ Leaves distinctly serrate.
= Pappus none : Mexico to Venezuela.

11. G. PATENS, Gray. Smoothish : stems flexuous, reclining,
becoming 5 m. in length, at length soft-woody at the base, striate-
angidate, sparingly and minutely appressed-pubescent ; leaves ovate,
acuminate, serrate, thin, obliquely truncate at the base, finely pubes-
cent, more or less scabrous above; petioles slender, 1 to 3.2 cm.
long; peduncles slender, finely appressed-pubescent; involucre
cylindric, 5 mm. in diameter; outer scales lanceolate acute or
attenuate, the inner longer, oblong, striate, obtusish : ligules 6 to
10, 1 cm. long, yellow turning whitish during or after drying; disk-
fiowers, 80 to 35. — Proc. Amer. acad., 5, 182; Hemsl., Biol. Cent.-
Amer. Bot., 2, 103. ? Wedclia cordata. Hook, and Arn., Bot.
Beech., 435. 3Iicroceph(dum ehrtnhergiamnn, Sch. Bip., and
Gymnolomia ehrenhergiiuia, Klatt, Leopoldina, 23, 90. Montanoa
thamasiiy Klatt, Abh. naturf. gesell. Halle, 15, 328. — State of
San Luis Potosi, Palmer, no. 1099, Pringle, no. 3937; Orizaba,
Schaff)ter, Botteri, no. 497, 810, Thomas\ Wartenberg in Hua-
steca, Eri^endherg, no. 95 ; Oaxaca, El Paridnetla, Comatii and
Ganzdlez, no. 906; Guatemala, von Tnerckheim, no. 287.

var. ABBREVIATA, Robiuson and Greenman. Stems 1 to 2 m.
long, sparingly appressed-pubescent: petioles very short, 4 to 6 mm.
in length ; leaves somewhat narrower and thicker than in the type :
pappus none. — Proc. Amer. acad., 29, 387. — Jalisco at Tequila,
Pr ingle, no. 4595 ; Guerrero, between Chilapa and Tixtla, altitude
1,600 to 2,100 m., E, W. Nelson, no. 2164.

var. gnatemalensis. With copious spreading pubescence on
the stem : pedicels 1 to 2.5 cm. long: pappus none. — Guatemala,

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ROBINSON & GREENMAN : CONTRIB. GRAY HERBARIUM. 95

San Miguel Uspantdn, Dept. Quiche, Heyde and Lux, no. 3870 of
Mr. J. Donnell Smith's sets.

var. brachypoda. Pubescence, at least on the upper part of
the stem, spreading : pedicels very short (2 to 5 mm. in length) :
pappus none. — Guatemala, Alta Verapaz, altitude 1,300 m., von
Tuerckheim, no. 852 of Mr. J. Donnell Smith's sets ; also in Colima,
S. W. Mexico, Palmer, no. 1223 (coll. of 1891).

▼ar. macrophylla. Pubescence of the stem copious, spread-
ing: leaves (1.5 dm. long, 8 to 10 cm. broad) much larger than in
the other forms of the species, subcrenately serrate : pappus none.
— Venezuela near To var, at 1,000 m. altitude, Fendler^ no. 686.

12. G. cosTARicENsis, Bcuth. Habit, foliage, and pubescence
as in G. patens : heads larger (disk-flowers about 50) ; involucre
campanulate : pubescence fine, appressed or subappressed : rays
about 10: achenes of the disk-flowers destitute of pappus but
sometimes crowned with an obscure ring or rim. — Benth. ace. to
Klatt, Bull. soc. hot. Belg., 81, 199, at least as to pi. of Oersted
(Pitticr's no. 2614, referred to this species by Klatt, is, in herb.
Klatt, jBaltimora scolosperum, Steetz!). G. rudbeckioidts, Hemsl.,
Biol. Cent.-Amer. Bot., 2, 163, in part, probably not HBK. Gym-
nopsisf costaricensis, Benth. in Oerst., Vidensk. meddel., 1852,
p. 90. G. vulcanica^ Steetz in Seem. Bot. herald, 157. Aspiiia
costaricensis^ Klatt, I.e., 201, as to synon. and Pittier's no. 8276
in part. — Costa Rica, Aguacate, and Ujaras, Oersted ; Rodeo de
Pacaca, Pittier^ no. 3276 in part, San Francisco de Guadalupe,
Tonduz^ nos. 7251, 7288; Rio Maria near S. Jos6, Tonduz, no.
7271 ; Panama Boquete, Veraguas, Seemaiin, no. 158S.

= = Pappus none : Peruvian.

13. G. RUDBECKioiDEs, HBK. Of habit and foliage of the
last two species but with short petioles (4 to 6 mm. long) , spread-
ing pubescence, and (ex icon.) more axillary or irregular (less
definitely corymbose) inflorescence. — Nov. gen. et spec, 4, 219,
t. 374. — Temperate regions, Agavaca, Peru, Humboldt and Bon-
plaTul. Not seen by the writers, who have been unable to match
satisfactorily the original description and figure of this species with
any Mexican or Central American plant.

= = = A delinite pappus present, consisting of several fimbriate scales.

14. ? G. suBPLExuosA, Benth. and Hook. f. Habit, foliage, pu-
bescence, and inflorescence much as in G. patens^ from which we are

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96 PROCEEDINGS : BOS'iX)N SOCIETY NATURAL HISTORY.

able to separate it solely by the pappus. — Gen., 2, 364 ; Hemsl.,1. c,
163. Wedelia subjfexuosa, Hook, and Arn., Bot. Beech., 435. —
Realejo, Nicaragua, Sinclair; Orizaba, Botteri, no. 435 (ace. to
Hemsl.). To this species of which we have seen but a single
achene we refer with doubt 8j)ecimens from the following localities :
Vera Cruz, Valley of Cordova, Bouryeau^ no. 1961; Oaxaca, Jaya-
catlan, Z. C, Smith, no. 896, Monte Alban, Pringle, no. 4859,
C. Z. Smith, no. 244, Valley of Etla, Z. C. Smith, no. 849,
Alvarez^ no. 723 ; Costa Kica, Prov. of San Jos^, */. Doimell
Smith, no. 4871. A better knowledge of the type of this species
will doubtless justify at least a varietal separation of some of the
specimens here mentioned.

•^ ^ Leaves entire or nearly so, canesceut -sericeous beneath.

15. G. f'AXEscENs, Robinson. Leaves oj)posite, ovate, subtrun-
cate or subcordate at the base, acute or often obtuse at the apex,
finely gray pubescent above, silky and silvery beneath ; petioles
2 cm. long: branches of the naked inflorescence long, bearing few
clustered heads at their ends. — Proc. Amer. acad., 27, 174. — San
Luis Potosi, brackish marsh, Las Tablas, Pringle, no. 3611, alkaline
plains. Hacienda de Angosture, Primjle, no. 3763.

•^ -^ Heads somewhat larger (of medium size) rather few, mostly solitary or
borne by 3's at the ends of the branches.

•»- Outer bracts of the involucre shorter than or little exceeding the inner :
species of Mexico, Cent. America, and Andean South America.

= Leaves sessile or petioles short (2 to 8 mm. long).

a. Pappus none : species of Mexico.

16. G. ovATA, Gray. Stem 4 to 8 dm. long or more, covered
with a short spreading pubescence : leaves ovate, subsessile, 3 to 7
cm. long, 2 to 4 cm. broad, serrate, mostly acute: heads 3 to 8,
long, unequally long-peduncled, 1.5 to 1.8 cm. in diameter (exclu-
sive of rays), many- flowered ; the bracts of the involucre narrow,
loose, very numerous. — Proc. Amer. acad., 19, 4. — Chiapas, Ghies-
breght, no. 554 ; Oaxaca, Sierra de San Filipe, altitude 1,800 to 2,500
m., Pringle^ no. 5674, Conzatti and Gonzdlez, no. 551.

17. G. LiEBMANXii, Klatt. Smaller: leaves sessile, cordate,
green on both sides, 2 cm. long, nearly as broad : heads few, 1 or 2
or perhaps 3 at the ends of the branches ; involucres 6 mm. in
diameter ; bracts 2-seriate, rather few, lanceolate ; rays about 9, con-
spicuously 5-nerved. — Leopoldina, 23, 90. Microcephalum lieb-

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ROBINSON & GREENMAN : CONTRIB. GRAY HERBARIUM. 97

mannii, Sch. Bip^ ace. to Klatt, 1. c. — S. Andres, Mecatlan, Mex-
ico, Liebmann, no. 588. Type in the herbarium of the Botanic
garden of Copenhagen ; a tracing and fragment in herb. Gray.

6. A rudimentary pappus generally present a« a short-fringed cup : South
American species.

18. G. TEXELLA, HBK. Prostrate, much-branched: stems slen-
der, hard : leaves opposite, ovate, subcordate, obtusish, thick, rugose,
very scabrous above, short- j)etioled, serrate or seiTulate, 2 to 5 cm.
long : heads solitary, terminal, on filiform peduncles ; involucre 8
to 10 mm. in diameter; the bracts oblong-lanceolate, canescent-
pubescent; disk achenes obovate, apj^ressed-pubescent, somewhat
contracted above into the at length obscure pappus-cup. — Nov.
gen. et spec, 4, 21H, t. 878; Klatt in Engl. Jahrb., 8, 42. —New
Granada near Ibague and Contreras, lhnnholdt\ in bushy under-
growths and on savannas of the Kio Dagua, Cauca, altitude 1,000
to 1,S00 m., Lthmann, nos. 19r)(), 2X65, 2994.

19. G. iiONDENsis, HBK., I. c, 21S. From description close to
if not identical with the preceding: leaves said to be subacumi-
nate at the apex and rounded at the base. — Torrid regions of New
Grenada near Honda, Humboldt, Sprengel (Syst., 3, 610) reduces
this species to G. tenella^ where it may well belong.

20. G. TRiPLiNERviA, HBK. Branches terete, smooth, glabrate,
the younger with appressed pilosity: leaves opposite, ovate, nar-
rowly subacuminate, acute at the base, crenate-dentate, reticulate-
veiny, 3-nerved, hispidulous with appressed hairs on both surfaces,
about 6 cm. long, half as broad; petioles 6 to 8 mm. long: heads
solitary on long appressed-pubescent peduncles; involucral bracts
numerous, appressed-pubescent, the outer slightly exceeding the
inner: disk-flowers and achenes unknown. — Nov. gen. et spec,
4, 219. — On high plains of Bogota, U. S. Colombia, Ilumholdt.
Not seen by the writers, the descr. compiled.

= = Petioles longer (8 to 20 nun. in leninh): ligules rather showy, 1 to 2
cm. long.

a Rays about 12, narrow, strongly carinate or conduplicate.

21. G. CRUCIATA, Klatt. Tall erect shrub, 3 m. high : branches
tetragonal, rough : leaves ovate- oblong, shortly cuneate or one-
sided at the base, 7 cm. in length, half as broad ; the petioles 1 cm.
long : rays about 12. — Bull. Herb. Boiss., 4, 480. — Near Castanal,
U. S. Colombia, Sonntag^ no. 8, June, 1888. Type in herb. Univ.
of Zurich ; a head and sketch in herb. Gray.

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98 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

6. Rays 8 to 10, not conspicuously cariuate or couduplicate : South Americau.

22. G. QUiTENsis, Bentb. and Hook. f. Shrub witb rather
copious pubescence : leaves all opposite, acuminate, from subcordate
to shortly cuneate at the base, 4 to 9 cm. long, half as broad
heads cymose, the peduncles rather short, 2 to 2.5 cm. long
much exceeded by the leaves. — Gen., 2, 364. Andrieuxia qxdten
sis, Benth., PI. Hartw., 206. — Andes of Ecuador, Hartxceg, no,
1142, Hall, Couthouy,

■^ •*^ Outer bracts of the involucre elongated, the inner short : species of Cen-
tral Brazil.

23. G. KUNTHiANA, Baker. Erect perennial pubescent herb:
leaves petiolate, ovate, serrate, the upper alternate : involucre 1 to
1.2 cm. in diameter; outer bracts oblong, large, unequal, acute, 1.2
to 2.4 cm. long: ligules 10 to 15. — Baker in Mart. Fl. Bras., 6,
pt. 3, 172. Gyynnapsis kwithiana^ Gardner in Hook. Lond. journ.
bot., 7, 292. — Dry mountainous regions near Concei^Ao Goyaz,
Brazil, Gardner, no. 3846.

* * * Heads and involucres as in the last group : leaves (all opposite) narrowly
lanceolate : pappus none : South American.

•.- Pedicels bracteolate.

24. G. GOEBELii, Klatt. Finely and cinereously appressed-
pubescent : leaves rather small : branches terminated by 3-headed
cymes; pedicels long and slender, bearing above the middle two
subulate bractlets; heads 1.2 to 1.5 cm. broad exclusive of the 16
to 20 narrow linear rays. — Klatt in Goebel, Pflanzenbiologische
schilderungen, 2, 49. — Dry slopes near the small town of Muru-
chies, Venezuela, K. GoeheL

■»- -.- Pedicels naked.

25. G. HiRsuTA, Klatt. Branching shrub, scarcely or not at all
canescent : leaves long-lanceolate, caudate-attenuate, obtusish at the
base, 7 cm. long, 1.5 cm. broad, bullate-rugose above, veiny and
pubescent beneath : ligules about 12, oblong, 5 mm. broad. — Klatt
in Engl. Jahrb., 8, 42 (18^<6). — Among shrubs, borders of woods,
Tungu-ragua, Ecuador, altitude 2,200 m., Lehmann, October, 1879,
no. 360 a.

**** Ht^ads laree, but liuules short; involucral scales oblong, the tips
slightly squarrose ; peduncles long : leaves oblong or elliptic, sessile, harsh
in texture and hirsute : perennial herb.

26. G. PARKiNsoxii, Hemsl. Erect, coarsely pubescent and

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ROBINSON & GREENMAN : CONTRIB. GRAY HERBARIUM. 1K>

scabrous : leaves alternate or scattered (the middle ones occasion-
ally verticillate) , 6 to 13 cm. long, 1/2 to 3.4 cm. broad, promi-
nently 3-nerved : beads on long stout ascending alternate peduncles,
subglobose, 2 to 2.5 cm. in diameter. — Biol. Cent.-Amer. Bot.,
2, 163. G. rudisy Gray, Proc. Amer. acad., 22, 424. G. rudisy
var. minor^ Rob. and Greenm., ibid, 29, 3S7 (small form) . — The
type (in herb. Kew), collected in Mexic/O, without locality, by Park-
inson, has brownish-purple flowers. This form is represented also
by Palmer's no. 531 from the Rio Blanco, Jalisco, Pringle's no.
4584 from rocky hills near Tequila (small form) , and Pose's no.
3031, coll. between Bolanos and Guadalajara, and no. 3700 coll.
at Bolanos. This typical form is connected by variegated transi-
tions with a forma flaviflora in which both disk- and ray-flowers
are golden yellow. The latter form is rej)resented by Pringle's
DO. 2460, from hillsides near Guadalajara, Pabner's no. 533 from
the Rio Blanco, and Pose's no. 2827, coll. between Colotlan and
Bolanos, Jalisco. No morphological differences between the indi-
viduals with brown-purple flowers and those with yellow flowers
have been detected.

* • • • « Heads lar;^ (1.6 to 1.8 cm. broad excl. of rays), terminal, solitary- ;
involueral bractn narrow, lance-linear, attenuate: rays long: leaves lance-
oblong.

27. O. serrata. Perennial : stems several, 5 dm. high, un-
branched, striate, terete, puberulent, leafy, 1 (-2)- headed: leaves
lance-oblong, opposite or alternate, attenuate, conspicuously ser-
rate, shortly petiolate, scabrous- puberulent above, scarcely paler,
pubescent, and reticulate veiny beneath, 6 to 12 cm. long, 1 to 2
cm. broad : involueral bracts herbaceous, lance-linear, attenuate,
appressed-canescent-pubescent especially on the mid-nerve, 1 to 1.2
cm. long: rays deep yellow, oblong, 1.5 to 2.5 cm. long, 5 to 8
mm. broad : disk-corollas yellow, essentially glabrous, i\ mm. long :
immature achenes 4 mm. long, glabrous, destitute of pappus. —
Collected in S. W. Chihuahua on Mt. Mohinora, 1 September, 189s,
E. W. Nelson, no. 4891. Types in herb. U. S. nat. museum and
herb. Gray.

• •«««« Heads of medium size, most nf them sessile or shortly pecfuncled
in few many-headed leafy corj-mbs : involueral bracts ovate, obtuse, con-
spicuously squarrose : leaves ovate, sessile.

28. G. SQUARROSA, Bcuth. and Hook. f. Erect perennial herb,
simple below, coarsely and copiously pubescent: leaves serrate

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100 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

scabrous above, paler and somewhat canescent-pubescent beneath,
5 to 7 cm. long : ligules (about 13) 1 to 1.3 cm. long. — Gen., 2, 3G2 ;
Hemsl., 1. c. ; Gray in Wats., Proc. Amer. acad., 22, 424. Zaliiza-
nia squarrosa^ Sch. Bip., Flora, 1S64, p. 217. — Ravines near Guada-
lajara, Jalisco, Oliva (lHb4) , Palmer, nos. 486, 741 (1886), Pm?^/c,
nos. 2194, 2474 (1888-89) . Flowering from September to Novem-
ber.

«««*««« Heads small or iiR'diuni -sized, s lender-ped uncled : involucral
bracts (all but a few (»f the outermost) broad, obovate or oblon«r, mostly
obtuse or rounded at the apex : leaves ovate or lanceolate to linear-oblon*;.

+- Scales of the involucre not canescent.

29. G. LATiBRACTEATA, Hcmsl. Stems terete, striate, purple,
alternately branched above, sparingly scabrous-pubescent: leaves
alternate (so far as known), lanceolate, attenuate at each end,
short-petioled, obsoletely serrate, 6 to S ("to 18") cm. long, 1 to 3
cm. broad : peduncles o to 7 cm. long, 1-headed; heads (excl. of
ligules), 1.5 cm. in diameter; involucre campanulate: rays few, 1.2
to 1.4 cm. long. — Hemsl., 1. c, 162. — Cerro Pinar, North Mexico,
Stemann, no. 1485. Collected but once.

-•- -4- Scales of the involucre canescent.

30. G. EXSiFOLiA, Benth. and Hook. f. Stems slender, terete,
strigillose, and scabrous: leaves opposite, sword-shaped, 1 to 1.2
dm. long, 1 to 1.5 cm. broad, gradually narrowed from near the
sessile base to the attenuate apex, very scabrous above, pale beneath :
heads cymose ; involucral scales all rounded at the apex, the outer
much shorter. — Gen., 2, 364; Hemsl., 1. c, 161. Montac/)iea ensi-
folia, Sch. Bip. in Seem. Bot. herald, 304. 2kiluzama eitsifolia,
Sch. Bip., Flora, 1864, p. 216. — Sierra Madre, Northern Mexico,
Seemann, no. 2007 in part; Tepic, Palmer (coll. of 1892).

31. G. SERICEA, Klatt. " Suffrutescent " : leaves chiefly oppo-
site, ovate, shortly connate-petiolate, entire, 3-nerved, acute, acu-
minate, 3 to 4 cm. long, half as broad, above green and scabrous,
beneath silky canescent: heads 1.2 cm. in diameter; involucral
bracts biseriate, oblong, subequal : rays about 20. narrowly oblong.
— L^opoldina, 23, 90. Mia*ocephalum sericetim^ Sch. Bip. ace. to
Klatt, I.e. — Mexico, without locality, JJebma7i7i, no. 609. Type
in herb. Botanic garden of Copenhagen; excellent drawing and
some fragments in herb. Gray.

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ROBINSON & GREENMAN : CONTRIB. GRAY HERBARIUM. 101

««»««••« Heads large (1.7 to 7 cm. broad excl. of rays) ; involucral bracts
broad, the inner large, spatulate with rounded apex: perennial herbs:
leaves lanceolate to oblong or ovate ; petioles sliort winged, or almost none.

1- Leaves oblong-lanceolate, pale beneath.

32. G. GHiESBREGHTii, Hemsl. Erect, simple or branched from
the base, 6 to 15 dm. high: leaves all alternate or some opposite,
serrate or serrulate, mostly acute at each end, 5 to 10 cm. long, 1
to 4 cm. broad: heads few, 2 to 2.5 cm. in diameter; outer scales
short, squarrose: ligules about 20, golden yellow, 2 cm. long. —
Hemsl., 1. c, 162. — South 3Iexico, hills of Oaxaca, Ghiesbreght,
nos. 385 (ace. to Hemsl.), 3S2, Printjle^ no. 4903, E. W, Kelson y
no3. 1371, 1464, L, t\ Smith, no. 799, Coiizatti and Gonzdlez^ no.
553 ; Michoacan on wooded hills near Patzcuaro, Fringle, no.
3339.

1- ■»- leaves elliptical, green beneath.

33. G. DEcuMBExs, Robiuson. Stems several from a lignescent
stock, decumbent, 4 to 5 dm. long: leaves chiefly opposite, 2.5 to
4 cm. long, half as broad, scabrous upon both surfaces: heads
sohtary or borne by threes at the summit of each stem ; outer scales
of the involucre ovate-lanceolate, acute, not squarrose, dark colored,
covered with appressed white hairs. — Proc. Amer. acad., 26, 165. —
Rocky hills, Tultenango, State of Mexico, Pi-inyle^ no. 3263 in
part; Zacatecas between Bolanos and (Guadalajara, Bose^ no. 3044
(leaves somewhat longer).

The achenes of this species are perfectly glabrous, obovate-
oblong, strongly compressed, striate, and completely destitute of
pappus. It is now found that another plant of astonishing similarity
was also collected and distributed under the type number (Pringle's
no. 3263). While in all observed external characters it cannot be
distinguished from the former, it has thick obtusely 4-angled mot-
tled and pubescent achenes which are crowned with 2 (to 4) palea-
ceous awns and intermediate scales. The latter plant seems to be
identical with Viguiera ghiesbreghtiij Gray.

-!-■»-••- Leaves ovate, large and broad.

•M. Petioles not auricled.

= Outer involucral bracts large, squarrosely spreading or reflexed,
about as long as the inner ones, acute or acuminate.

34. G. FLAVA, Hemsl. Stems strigillose becoming hirsute with
coarse spreading hairs toward the summit and upon the peduncles :

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102 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

leaves 7 to 10 cm. long, 2 to 3 cm. broad, above scabrous, beneath
hirsute upon the veins and finely canescent-pubescent upon the
surface: heads (excl. the rays) '2.5 to 3.5 cm. in diameter: ligules
about 15. — Hemsl., 1. c, 161. — Oaxaca, Ghiesbreght, no. 216
(type in herb. Kew). We follow Dr. Gray in referring to this
species GhiesbreghVs no. 3^<8, from which the above characteriza-
tion is drawn.

35. G*. megacephala. Stems simple, striate, scarcely at all sca-
brous, covered with minute appressed hairs, also lanate in lines
decurrent each from either side of the short winged petioles : leaves
ovate-lanceolate with short slender caudate acumination, serrate,
scabrous above, hirsutulous upon the veins, but green and glabrous
upon the surface beneath, 8 to 12 cm. .long, 2 to 4.5 cm. broad:
heads terminal, solitary, about 7 cm. broad : no rays observed. —
Collected between Tlapa and Ayusinapa, Guerrero, Mexico, alti-
tude 1,380 to 1,700 m., 13 December, 1894, E. W. Nelsoji, no.
2105, distributed as G.flava.

var. simulans. Pubescence of the stem coarser, spreading : bases
of the hairs enlarged, white, tuberculate : leaves somewhat larger,
less pubescent, merely cuspidate at the obtusish tip: rays about 13,
oblong, bright yellow, 3 cm. long, 7 to 10 mm. broad. — Collected
on the Sierra de los Morones near Flateado, Zacatecas, altitude
2,300 m. 1 September, 1897, Dr. J, N, Rose, no. 2740. This vari-
ety closely simulates Viguiera excelsa, Benth. and Hook.f., which,
however, differs technically and, as it appears, with constancy in
its pubescent achenes which bear at the summit two conspicuous
awns and some intermediate scales.

= = Outer involucral bracts much shorter than the large inner ones.

36. G. PLATYLEPis, Gray. Hirsute shrub, 1 to 3 m. high : leaves
chiefly opposite, acuminate or attenuate at each end, very sca-
brous above, paler and densely soft-pubescent beneath, 1 dm. long,
3-nerved from considerably above the base : peduncles thickened
upwards; heads (excl. of rays) 2 to 3 cm. in diameter; ligules
15 to 20 in number, 2 to 3 cm. long. — Proc. Amer. acad., 19, 5;
Klatt, Bull. soc. bot. Belg., 31, 199. G, decurrens^ Klatt, Leopol-
dina, 23, 90. Peri7nenio2ysi8 jyerfoliata^ Sch. Bip., ace. to Klatt,
1. c. Tithonia scaherrima, Benth. in Oerst., Vidensk. meddel.,
1852, p. 91. T, platylepis, Sch. Bip., ace. to Benth. and Hook.f.,
1. c, 368 (name only). JlirasoHa scaberrima, Benth. and Hook.f.

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ROBINSON & GREENMAN : CONTRIB. GRAY HERBARIUM. 103

ace. to Ilemsl., I. c, lOS. — South Mexico, Mirador, JAehintfmi^ no.
251 ; Chiapas, Ghieahreght^ no. 572, Caec. and Ed. Seler, no.
2187 ; Valley of Cordova, Bonrf/edu, no. 1<S51 ; Vera Cruz, J^rin-
gle^ no. 60H7 ; Orizaba, Grtn/ ; (4uateniala, ron Tnerck'hehn^ no.
354 of Mr. J. Donnell Smith's sets; Costa llica, Pitficr, nos. :il.*i6,
3735, Tonduz, no. 719().

•H* ^ Cuiieately winged petioles tendinis t(» eiihir^emeiit and usually auiicled

at the base.

37. (t. oalva, Gray. Shai^gy- pubescent with long white silky
hairs: liabit as in Tithoaia: leaves alternate, broadly ovate, acu-
minate, abruptly contracted to winged and (sometimes obsoletely)
auriculatc petioles: involucre hirsute. — Proc. Anier. acad., 11), 5;
Kook. f. and Jacks., Ind. Kew., 1, 107G. Tithouhi cnhui^ Sch. Kip.
in Seeraann, 1. c, 305. Mirasolia calro^ Benth. and Ilook. f., Gen.,
2, 368; Hemsl., 1. c, 168. Sierra Madre, Northern Mexico, fSee-
mann^ no. 2045 ; Durango, Ro8(\ no. 2293.

var. lancifolia. stem sparingly hirsute or villous : leaves
lanceolate. — Tepic, Aca])oneta, February, isy.% F. J[. Zr/W>, no.
539. Types in herb. Gray and herb. U. S. nat. museum.

Doubtful species.

G. coNNATA, Spreng., Syst. 3, 610 [conudtion). — Branches
hispid : leaves oblong, attenuate at either end, connate, hispidulous :
]>eduncles terminal, short: foliaceous involucre subsimple. — 13razil,
tSeUo, A species not mentioned in Martins, Fl. Hras., and probably
not of this genus. The description is a translation of the original
characterization .

G. MiCROCEPHALA, Lcss. Herb with 3-nerved leaves, small
cylindrical discoid short-pedicelled heads: leaves petiolate, opposite,
paler beneath, oblong-ovate, long-acuminate, very obtuse at the
base, sharply serrate, hirsute-hispid, scabrous, 3.1 to 5 cm. long, 1 to

3 cm. broad : scales of the involucre passing from Hat semilanceo-
late foliaceous bracts, hirsute on the midnerve, to longer (about

4 mm. long) and broader dry linear shortly acuminate glabrous
bracts, fimbriate-dentate at the a})ex, scarious on the margin, and
traversed by green nerves : branches opposite, diverging at the
apex into 3 short filiform branchlets bearing several hea<ls (4 mm.
long) above. — Linnaea, 5, 153, whence the foregoing description is

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104 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

translated. — Near Hacienda de la Lagiina, October, Schiede and
J}ej}pe\ also Cordillera of Oaxaca, (rakiftfi^ no. 2090 (ace. to
Ilemsl.). Exceptional in its discoid heads and perhaps not of this
genus.

Transferred or suppressed species.

G. decurrexs, Klatt, Leopoldina, 28, 90, is G, phttyhjjia, Gray,
— an identity recognized by Dr. Klatt himself as shown by a
manuscript note in his herbarium.

G. EHRENiiERciiAXA, Klatt, Lcopoldiua, 28, 90, is G. jjote/ts,
Gray.

G. RLTDis, Gray (in Wats., Proc. Amer. acad., 22, 424) is (r.
jxirkinsonii^ Hemsl., Biol. C'ent.-Amer. Bot., 2, 1(]8, — an identity
kindly verified by Mr. Ilemsley by comparison of the types.

G. siLVATK'A, Klatt, Leopoldina, 2r), 104 (1SS9); Bull, soc, bot.
Belg., 81, 199 (1S92). 1 Representing this species in the Klatt her-
barium we find two specimens collected by Lehmann at Cuenca,
altitude 2,000 m., 10 October, 1S79, Pittier's no. 09S7, collected at
San liafael de Carthago, altitude 1,5()0 m., 2S August, 1S92, and
an excellent drawing of a specimen collected at Mt. Irazu, Costa
Rica, by Dr. Hoffman. All of these appear to be identical and to
agree in all more conspicuous characters with the original descrip-
tion. Nevertheless, careful dissections show that the ray-achenes
are fertile and the ligules sessile indicating that the sj)ecies is a
Heliopsis.

G. TRiLoiJA, Gray, with fertile ray-fiowers and an unequal scale-
like pappus, is certainly a Zaluzania. The specific name triloba
having been used in the latter genus the species may be transferred
as Z. grayiana, n. comb.

G. TRiPMXERviA, Klatt, in Engl. Jahrb., 8, 42 (not IIIU^.), is
a Heliopsis, the ray-fiowers being distinctly jnstillate and fertile,
while the ligules are sessile, not contracted into a slender tube at
the base.

Gymnopsis veruesinoides, DC. (Prodr., 5, hi'A) of the Island of
Trinidad is an Aspilia.

Gymnopsis viLCANirA, Steetz in Seem. Bot. herald. If)?, appears
from the description and the examination of a single head to be
Gynuiolomui cost^wicousis^ Benth.

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ROBINSON & (iREENMAN : CONTRIB. GRAY HERBARIUM. 105

2. SUPPLEMENTARY NOTES UPON (^ALEA, TUIDAX,
AND MIKANIA.

Some years ago the writers (Proc. Araer. acad., 32, 1-30) piib-
lisheil revisions of the genera Ualea, Tridax, and Mikania. These
synopses were based chiefly upon the representation of the genera
in the Gray herbarium, and considerable difficulty was naturally
experienced in the exact placing of some of the species of European
writers whose types were not readily available for examination.
The recent acquisition of the Klatt collection by the (4ray herba-
rium does much to remove these difficulties and puts the writers
in a position to add the following corrections and supplementary
notes upon their former revision.

Galea peduncularks, var. lonmufolia, Gray. Dr. Rose's no.
2730, from the Sierra de los IVIoronea, near Plateado in the State
of Zacatecas, corresj)onds closely with Cnhd pedtmcularis, var.
longifolUt^ Gray, but has pap])us, thus showing that the achenes
of this variety are no more constant in regard to pappus than those
of the more ty[)ical forms of the species. The variety is, however,
readily distinguished by its long leaves.

C. PERFORATA, Klatt, Lcopoldiua, :iO, 95, hfvs immature heads
but is otherwise like Fendler's no. 637 (from Tovar, Venezuela)
which was relegated witliout question by Schultz Bipontinus to C,
solkl(igine(t, HBK. On comparing Dr. Klatt's type (coll. on the
Mayquetea River by Edw. Otto, no. 4f>0) with Kunth's description
and plate (IIBK., Nov. ^aw, et spec, 4, 295, t. 407) we can find no
significant difference.

C. DENsiFLORA, Klatt, Lcopoldiua, 20, 9(), is Af/eratum coinj-
zouies, L.

C. PELLuriDiyERViA, Klatt, Bull. soc. hot. Belg., 31, 207, is not
to be satisfactorily distinguished from C (wilfaris, DC.

C. AXILLARIS, DC. To this species should also be referred C.
prunifolla, Klatt, 1. c, 208. in part (as to Pittier's nos. 3219, 3095,
4520, 4938, and 7023), not IIBK. The distinctions between C.
iisrillftris and its variety urticnefoUa seem to be scarcely worthy of
recognition.

C. pittierl sp. n. Shrub with slender flexuous terete puberu-
lent or toward the end tomentulose branchis : loaves ovate to
elliptic, thickish, crenate- serrate, acutish, 4.5 to 7.5 cm. long, 2.5

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10<) PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

to 8.7 cm. broad, rugose and very scabrous above, veiny grayish-
tomentose and resinous-dotted beneath, 5-nerved from near the
obtuse or subcordate base ; i)etioles 0 to 9 nira. long, covered with
a fine fuscous pubescence : heads numerous, rather small, discoid ;
pedicels filiform, 1 to 1.2 cm. long, some fascicled In the upper axils
but most of them in regular hemispherical umbelliform leafy-bracted
clusters (4 to 5 cm. in diameter) at the ends of the branches ; invo-
lucre cylindric-ovoid, somewhat turbinate at the base, the outer
scales very short, linear-oblong to narrowly ovate, herbaceous,
tomentulose, the intermediate oblong, obtuse, often erose and squar-
rose at the tip, the inner lance-oblong, acute, bright yellow, peta-
loid : achenes narrowly fusiform, angled, 2.7 mm. long, callose at
the base, covered with short spreading gray hairs ; pappus-awns
about 20, narrow, attenuate, 4.() mm. long. — C, pruinfolia, Klatt,
Bull. soc. bot. Belg., :il, 2()S, in part, not HBK. — Costa Rica,
banks of the Kio Ceibo near Buenos Ayres, altitude 200 m., Janu-
ary, 1892, Prof. If. Plttler^ no. 4918, Chennn de la Caldera between
San Mateo and San Kamon, P. Biolleyy 25 January, 1892, no.
7015; also by Pt-of. Pittier in woods at Boruca, altitude 450 m.,
February, 1S91. C. prunifoUn, HBK., to which this plant was
referred by Dr. Klatt, has larger leaves nearly or quite smooth and
somewhat coriaceous. It also differs in the calyculate bractlets of
the involucre, which are suborbicular and of larger size.

C. GRAYii, Klatt, Leopoldina, 20, 90, overlooked in our revision,
should replace C. to)nentosa^ Gray, not Gardner.

C. TKKNiFoLiA, Oliver, Trans. Linn. soc. ser. 2, 2, 277, t. 48, figs.
9-10 (188()). For this species the name of which is antedated by
n. ternifoU'i, IIBK., Nov. gen. et spec, 4, 294, we would propose
C. oliverii.

TiUDAX (iALEOTTii, Klatt, 1. c, at least as to the plant with lobed
pilose leaves. T. tnhtrosa, Rob. and Greenm., Proc. Amer. acad.,
82,4. In the Klatt herbarium T. fftth ottii is represented by two
sheets of well executed drawings. On one is an excellent repre-
sentation of the j)lant we have called T. tuberosd.^ with lobed leaves,
cuneate at the base and distinctly hirsute. On the other sheet two
])lants are represented, one being identical with that just mentioned,
the other having the more oval serrate unlobed leaves of T. brachy-
Ifpf's^ llemsl. Dr. Klatt 's description clearly shows that he had in
mind the plant with lobed leaves. The description, however, is quite
in error as to the involucral scales being 2-seriate. They are clearly

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ROBINSON & GREENMAN : CONTRIB. GRAY HERBARIUM. 107

3— 4-8eriate in both his drawings and in a single autlientic head pre-
served in a pooket on one of the sheets of drawings. In the
arrangement of species T. galeottii should, accordingly, be inserted
where T, tuber osa has been placed.

T. iMBRicATA, Sch. Bip. in Klatt, Flora, 1885, p. 202. T. petro-
phitot Rob. and Greenm. 1. c, 5. The authentic specimen of T.
inibricata in the Klatt herbarium is very poor, but sufficient to
show with a fair degree of certainty the identity of T. 2>etrophila.
The involucre is here also 8-4-8eriate, not biseriate as we had
supposed.

T. vEETiciLLATA, Klatt, Lcopoldiua, 25, 107 (1889). This
South American species, omitted from our revision, is the only
member of the genus with verticillate leaves. In the arrangement
of the species it may well be placed in subgenus Eutridax after T.
angtistifolia,

T. BHBBNBEBGii, Sch. Bip. in Klatt, 1. c. Klatt's drawing of
this species shows a plant with long internodes and habit of T,
dubia^ but the achenes preserved in the Klatt herbarium have the
pappus rather of a Galea than a Tridax. The species must remain
doubtful until more complete material can be seen.

MiKANiA OLIVACKA, Klatt, Bull. soc. l)ot. Holg., 31, 11)5. — An
examination of the type in the Klatt herbarium shows it to be
Pittier's no. 4938, not 4433, as cited in the original publication of
the species.

M. PUNCTATA, Klatt, 1. c, originally described as having the
heads 7-flowered, proves, on careful dissections of portions of the
type material, to have uniformly 4 flowers in each head, fully
agreeing in this regard with the other sj)ecie8 of the genus.

M. FENDLERi, Klatt, Abh. naturf. gesellsch. Halle, 15, 324,
although reduced in our revision to a synonym of 3f. cordlfoUa^
Willd., is probably worthy of specific rank.

M. gonzalezii, sp. n. Vigorous smoothish twiner : branches
obsoletely 6-angled, finely striate, glabrous : leaves large, thin, gla-
brous, ovate, entire, acute, broadly and shal lowly cordate or sul>-
cordate, 5-nerved from the base, 4 to 12 cm. l')ng, 3 to 10 cm.
broad; petioles 1.5 to 5 cm. long: heads in an elongated open
panicle with short opposite corymbiferous branches (5 to 15 cm.
long); bracts of the inflorescence oblong-lanceolate, ])etiolate ;
involucres 7 mm. long, subtended by shorter sessile oblong-ellip-
tic or oval bractlets ; pedicels puberulent under a lens; involucral

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108 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

scales oblong, essentially glabrous except at the obtusish or barely
acute tip : corolla 5 to 6 mm. long, about equalling the sordid pap-
pus : achenes 4 mm. long, nearly glabrous. — Collected by Prof.
C. Cofizatti at Colonia Melchor Ocampo, Vera Cruz, Mexico, alti-
tude 1,200 m., 8 December, 1S95, no. 18, and by Prof. C. Comatti,
and V. Gonzdlez in Canton de Cordoba, Vera Cruz, 27 December,
1897, no. 637 in part. This noteworthy plant is clearly distinct
from any Mexican species known to us, nor have we been success-
ful in referring it to any South -American species. In the genus it
should, probably, be placed near M. cordifolut. from which, how-
ever, it differs in the size, form, texture, and glabrous character of
the leaves, which are more or less nigrescent in dr^nng. We take
pleasure in dedicating the species to Mr. V. Gonzdlez, who as the
able assistant of Professor Conzatti, has done much to further the
knowledge of Mexican plants.

Printed, Awjud, ISOO.

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Proo#«diitg» oi the BostDti Socittty ol Ifatixred History.
Vou 29, No. f>,

TJIE DEVEUJPMEJrr OF PEMl-IA St IIMACKERI lUrHAHD.

By Mkkviw T, ^itiJLKii*

Wrrii fHict;u rr*\TK!*.

BOSTON:
PKINTED FOR THE SOCIETY.

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No. 6. — Ths Development of Penilia schmackeri Richard.
By Mervin T. Sudler, Baltimore, Md.
With three plates.

During June, 1896, immense swarms of small crustaceans sud-
denly appeared in the water of the harbor of Beaufort, N. C.
Their 'numbers were so great that the animals usually found near
the surface of the water were completely obscured and, in towing,
the meshes of the net after being drawn through the water a few
yards were clogged with these crustaceans. This abundance
lasted but a few days when the creatures disappeared completely
and as suddenly as they had appeared. Dr. C. P. Sigerfoos, then
of the marine laboratory of the Johns Hopkins university, pre-
served a number for future study.

Three methods were used to preserve and fix these animals';
viz. : —

1. A saturated solution of corrosive sublimate used cold for a
few minutes.

2. A corrosive sublimate and acetic acid reagent, composed of
Corrosive sublimate (saturated solution) . . 90 parts
Glacial acetic acid 10 "

3. 80% alcoholic picro-sulphuric acid.

The animals were washed in water after treatment with each of
these solutions, run up through the graded series of alcohol until
they were jn 80%, and kept until used. The material so preserved
was placed in my hands and was studied by means of sections of
the adults containing young and the separate embryos after they
had been removed and by means of adults and embryos mounted
whole. Kleinenberg*s haematoxylin as a staining agent gave the
best results in the first case, and where they were mounted whole a
few moments staining in Czokor's alum cochineal proved the most
satisfactory. Xylol was used as a clearing agent and Canada balsam
as a mounting medium.

Systematic Position.

The crustaceans were found to be Cladocera (Daphnoidea) and
belong to Penilia, of which two species have been described by

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110 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

Dana. All of the members of the Sididae thus far known are
marine and especially distinguished from all other families of the
Cladocera by the presence of six pairs of thoracic appendages and
by the fact that one or both of the branches of the second antennae
have less than three joints, the characteristic number for most of
the members of this group. The species has been determined by
Prof. E. A. Birge of Madison, Wis., who has compared the form
with the type specimens of M. Richard.

The embryology of no member of the Sididae has been studied,
and most of the work on the entire group has been of a systematic
or anatomical character. Only three papers of importance dealing
with the embryology of the Cladocera are known to me ; these are
by Grobben (79), Lebedinsky ('91), and Samassa ('98).

Other articles, such as those of Dohrn ('(39) and of Zaddach
('41), are old and give but few details of development.

Anatomy.

Penilia when alive is transparent and has no pigment of any kind
except the very small amount in its single median eye. It measures
from 0.8 to 1.0 mm. in length, not including the biramose swim-
ming antennae, which with their setae are about 0.8 mm. The
long pair of abdominal setae are 0.4 mm. in length, making the
entire animal from 2.0 to 2.2 mm. long. In general shape (Fig. 1)
it somewhat resembles Daphnia similis (Claus). At one stage
of the life of Penilia eleven pairs of appendages appear, but in the
adult ten only arc present. The small anterior, or first, antennae
are situated on tlie posterior side of the two prominent horns which
project downward from either side of the head. They contain
nerve celln, and one long sensory seta is attached to the anterior
side of their distal end and to tlie ]>ost(*ri()r side a pencil of small
sensory liairs. Tlie large biranio.se swimming antenna is composed
of one large basal joint from which extend an exopodite and endo-
j)odite, eaoh composed of two segments, the first about three times
as long as the secon<l ynui. These joints liave long, stiff setae to
ai<l in swinmiing. The mandible is strongly developed and moved
by comparatively large nniscles. The fust maxilla is composed of
a single joint. It is situated directly behind the mandible and is
much weaker than that appendage. It possesses setae on its end.

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SUDLER: DEVELOPMENT OF PENILIA. Ill

The second maxilla disappears quite early in the life history of the
animal, and takes no part in the anatomy of the adult. The six
pairs of thoracic appendages are all much alike and are all
biramose.

The heart is oval in shape and lies above the intestine just in
front of the brood-chamber and directly over the shell-gland. It is
composed of about thirty cells and has one pair of ostia situated
near the posterior end of the organ.

The digestive tract has no digestive pouch or gland, but a number
of gland cells around the pharynx may to some extent take its place
by secreting similar substances. The intestine curves over against
the dorsal side of the animal and terminates at the anus between
two long setae. The shell-glands are nearly round and are com-
posed of from six to twelve large cells surrounding a central lumen.
This lumen empties to the exterior in the angle formed by the shell
and the body of the animal. The shape of these glands in Penilia
is unlike that of any other crustacean. In their histological struc-
ture the cells show an outer striated deeper staining border and an
inner striated but lighter staining portion. The nuclei stain very
dark. These cells resemble in structure the secreting cells of the
homologous glands of AsUicvs fluviatilis figured by Grobben, but
they are much larger. This comparison must not be carried too far
as the proportion between the different staining portions is not alike.
Ordinarily phyllopods show only striated cells, both borders being
very much alike.

The ovaries are paired and lie on either side of the digestive
tract near the posterior end of the animal. They do not exhi])it
the cells arranged in groups of four as Glaus describes for Jhiphnin
magna or Weismann for Leptodovn hi/all tia^hwi the etrgs are in all
probability formed in the same manner, /. c. from four cells. Tlu*
absorption and coalescence of the four into one must be quite rapi<l
as transition stages are hard to find.

The brood-chamber lies above and to either side of the difjjestive
tract. In cross-section it is heart-shaped with the apex of tln»
heart up toward the dorsal si<le and the de])reHsi<)n at the larger
end of the heart being occupied by the intestine. Its size varies
greatly, depending upon the number and size of the embryos it may
contain.

No males were found among the many hundreds of specimens
that I have examined.

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112 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

The Development of the Eggs.

The eggs of Penilia when laid are oval and much longer than
broad, measuring from 100 or 110 /a in length and 20 or 30 fj, in
breadth. Usually from four to six are deposited at one time,
although the number may range from one in young or small ani-
mals to eight in large well-developed adults. Generally the same
number is laid on either side but exceptions to this are found.
According to Grobben the brood-chamber of Moiiia rectirostris
may contain twenty-two eggs, and he has seen as many as thirty
well-developed embryos in that of Moiiia paradoxa.

The eg^ of Penilia contains but little yolk and possesses a small
deep staining nucleus surrounded by a circle of clearer protoplasm.
They have a vitelline membrane and never at any stage do they
possess more than the one. Lebedinsky found two in Duphuia
aimilis^ a chorion and vitelline membrane. Grobben always found
but the one (vitelline) in Moina. I think the q^q of Penilia repre-
sents the surviving cell of four homologous cells, and for these
reasons. The ovary is usually seen to be made up of cells arranged
in quite a definite manner. At the posterior end the cells are of
different sizes and arranged in no regular order. This is the ger-
minal epithelium. At the anterior part a row of cells alike in size
and appearance occurs. These, however, are not arranged in sets
of four so as to give the whole a bead-like structure, as is seen in
Daphnia and Leptodora, but they are always some multiple of four.
As the usual number of eggs found on a side is three, so the usual
number of cells seen in ripening ovaries is found to be twelve. The
smallest number of these cells when any at all are present is four.
The size of these four cells is slightly larger than that of the mature
6gg ; t>ut the size of the qqq^ corresj)onds very nearly to the size of
them all, and is therefore very much larger than any one cell. In a
few (three) specimens I have seen what I have taken to be the act
of fusion. Nuclei in partially broken states occurred in protoplasm
which seemed separated in places and slightly broken and irregular.
This fusion of the four cells must occur quite rapidly as transition
stages are so rare. In some cases, in eggs which had not seg-
mented, small fragments of material which stained as nuclear
material appeared. These were embedded in the substance of the
egg, and I interpreted them as fragments of the nuclei of the three
cells that lose their identity in the formation of the egg, I found

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SUDLER: DEVELOPMENT OF PENILIA. 113

nothing that I could call a polar-body. This structure is present in
Daphnia similis until the eight-celled stage according to Lebedinsky
and the sixteen-cell stage in Moina according to Grobben. These
obserrers did not see the formation of the polar-body, but state that
it is formed while the egg is in the ovary. From these descrip-
tions it seems to me that there is a possibility of its not being a
polar-body but a disintegrating nucleus of one of the ovarian cells
which has entered into the composition of the egg.

The Position of the Egg in the Brood-Ch amber.

The eggs when laid lie in the brood-chamber with their long axes
parallel to that of the mother where not more than two are laid
on either side. In large individuals the orientation may be the
same with a larger number of eggs ; but in most cases where the
number on a side is more than two, the eggs overlap so that the pos-
terior end of each egg is outside the anterior end of the egg directly
behind it. In some cases this position is not assumed until cleavage
has advanced to the four- or eight-cell stage, and in a few cases it
may not be assumed until the appearance of the appendages. Later
on the change in position is controlled by different factors, such as
the change in the shape of the embryo and the change in the shape
of the enlarging brood-chamber. These will be described later.

The embryo is nourished by means of the blood-plasma or albu-
men with which it is surrounded. Its increase in size is due
entirely to material obtained in this way. In the preserved speci-
mens this nourishing substance has coagulated and causes the
embryos to adhere quite firmly to the mother. In sections showing
a segmentation cavity we find a coagulum slightly stained which
probably comes from the same source.

Segmentation.

The first cleavage plane is transverse to the long axis of the ogg
and divides it into two separate and distinct cells, the anterior of
which is slightly larger than the posterior. (Fig. 7.)

The next plane of division is through the long axis of the egg
at right angles to the first and is almost parallel with a plane pass-
ing through the middle line of the adult. A four-celled embryo
is thus formed with the two anterior cells slightly larger than the
two posterior ones. (Fig. 8.)

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114 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

Sections of eggs throughout their development show through the
protoplasm a sharply defined and delicate reticulum with coarse
meshes. In the young stages and with high magnification, small
enlargements can be distinguished where the threads meet to form
the meshes.

The third plane of division is longitudinal but perpendicular to
the second, forming an eight-celled embryo in which the cells of
the anterior end are still slightly larger than those of the posterior
end. (Fig. 9.)

In the fourth stage of cleavage the eight cells have divided
transversely making sixteen cells. (Fig. 10.) Two planes of
division were nt'cessary to cause tliis. The eight anterior cells can
still be seen to be a very little larger tlian the eight posterior ones.
At this stage (Trobben has already distinguished a cell as the
mother-cell of the genital organs, another as the primitive ento-
derm cell and still others surrounding the "genitalzelle" which
are thought to give rise to the mesoderm. Thus at this early stage
these few cells foreshadow all of the germ layers as well as the sexual
cells of the adult. These cells are distinguished 1st by their peculiar
staining properties and 2d by difTerences in their action ; t. e.
time of division in reference to the other cells of the egg. Samassa
in his studies on the same animal finds the so-called " Grobben'sche
zelle" ("genitalzelle " of Grobben) and the entoderm present but
not distinguishable to the same extent as Grobben claims. He is
unable to distinguish the '' Grobben'sche zelle " after the original
one has divided into eight. He fails to find that it develops into
any particular organ of the adult. In Penilia I can find no cell
marked from its fellows by any peculiarity at this stage. The cells
vary somewhat as to size but apparently not in such a definite
manner as on that account to allow any particular one to be chosen
and its history followed.

In the fifth or thirty-two-celled stage regularity of cleavage
planes is hard to observe and probably does not occur with exact-
ness. From the arrangement of the cells, however, the planes
must be longitudinal again in their genera] direction. It is no
longer possible to distinguish any difference in the size of the cells
lying at the anterior end from those at the posterior. A cross-
section of this stage shows segmentation to be complete with a
large segmentation cavity containing no yolk. (Fig. 11.)

The sixth or sixty-four-celled stage still shows no sign of differen-

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SUDLER: DEVELOPMENT OF PENILIA. 115

tiation. All of the planes of division cease to be distinguishable
as such from this time. (Fig. 12.)

The seventh stage probably has about one hundred and twenty-
eight cells and shows the beginning of gastrulation. (Fig. 13.)

Gastrulation.

This process always t«akes place in a very definite part of the
egg with reference to its position in the mother.

No matter on which side of the intestine of the adult the ogg
lies, gastrulation always occurs at its outer posterior comer. An
exception to this rule was not observed in the scores of specimens
examined at this stage. A median horizontal longitudinal section
(Fig. 12) at this time shows projecting into the segmentation
cavity from the outer and posterior angle of the outer wall of the
egg an indefinite amount of protoplasm containing one or two
nuclei. At the particular point to which this is attached to the
outer wall one or rarely two cells have sunk below the surface
and are in the process of migrating into the interior. (Fig. 13.)
The nuclei and protoplasm shown in the interior have reached
this position by a similar process. In all probability regularity in
the planes of division has disappeared even before this time, biit
it certainly has now. The cells, however, continue to divide at
nearly the same time. An older embryo (Fig. 14) shows that both
the amount of protoplasm and number of nuclei in it in the seg-
mentation cavity have increased and a larger num])er of cells are
concerned in the inwandering and formation of the mouth of the
gastrula.

In Moina, gastrulation begins later than in Penilia, and apparently
the process is more rapid since more cells are concerned from the
beginning. In an older Penilia larva this indefinite protoplasmic
mass has become quite a definite rod running through the segmenta-
tion cavity and contains numerous nuclei. The gastrula mouth is
much deeper and enlarged. Xo differentiation except that noted
can be seen. Cell walls are now made out with great difticulty, if
at all, although the individual cells stand out as such very plainly.
Boundaries between them are marked off by lighter staining areas
of protoplasm. (Fig. 15.) At a corresponding or even earlier
stage, Moina, according to Grobben, already shows the beginning

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116 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

•of the nervous system. Samassa states that the embryo is more
tadvanced before such a differentiation can.be distinguished.

Size of Embhyos.

So far in Penilia the embryos have grown but little, the elements
composing them simply becoming smaller with each succeeding
stage ; but from this time on, increase in size of the whole is marked
and rapid. In this respect it agrees with the development of Moina.
The embryos and eggs from the same mother are all of the same
size and stage, as far as can be detected, throughout their develop-
ment. Embryos of like stages but from different mothers vary
somewhat in size; a younger stage is larger than a more advanced
one from another animal. Grobben found the same difference in
size present in the embryos of Moina and thought the cause was
traceable to variation in the size of the eggs when first laid by the
different mothers. This furnishes a partial explanation for the
phenomena seen in Penilia as the early segmentation stages of this
animal show the same differences in size, but never to the same
extent that the older embryos do. The amount and rapidity with
which the nutritive blood-plasma is supplied to the embryos also
vary in all probability, and this may be the principal cause in the
greater variation in size seen in the older embryos.

The Germ Layers.

The protoplasmic rod occupying the segmentation cavity is com-
posed almost entirely of the entoderm. The mesodenn originates
on either side of the mid-ventral line at the angles of the gastrula,
which at these particular points cannot be separated definitely into
entoderm and mesoderm. Certain cells, instead of growing up into
the center of the segmentation cavity, project out into these angles
and toward the ectoderm. As they multiply they increase in length
and width, but are only one-layered in thickness. They leave the
entoderm completely and cling to the inside of the ectoderm.
(Fig. 16.) These and their descendants form the mesoderm of the
adult animal. Samassa finds on the ventral side of Moina a mass of
tissue somewhat semilunar in cross-section. The central-lying cells
form themselves into a solid cylindrical row. These are the ento-

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SUDLER: DEVELOPMENT OF PENILIA. 117

derm, and the rest to either side of them are meso.derm. As has
been shown, the separation in Penilia is much earlier. The facts
observed in Penilia correspond to those given by Lebedinsky for
Daphnia similis, where the mesoderm spreads forward as two sym-
metrical rows from the gastrula invagination. At a later stage in
Penilia (Figs. 17-23) the mesoderm has grown slowly forward and
together at the mid-ventral line, making the outer wall two-layered
with the exception of the dorsal side where only ectoderm exists.

The gastrula mouth gradually becomes smaller, and by the time
the second antennae show it has disappeared almost completely,
leaving its position marked only by the entodermal rod, which soon
breaks away, making it impossible to locate the spot where the
anus breaks through. Grobben concludes that the mouth of the
adult Moina arises directly from the gastrula mouth of the larva.
This derivation of the mouth is impossible in Penilia, as will l)e
seen later. In fact, it is the rule throughout the Crustacea for the
anus and not the mouth to be formed from or near the position
occupied by the gastrula mouth.

The derivation of the anus in Penilia is open to question. The
anus appears in the middle line arid in the same or immediate
vicinity as that occupied before by the gastrula month. The anus
may be formed de novo slightly above or below this point. The
gastrula mouth completely disappears, and so it is impossible to say
whether it ever reopens.

The Appendages and Change in the Position of the

Embryos.

About this time a change in the outward form of the embryos
makes itself apparent. The gastrula mouth still lies at the posterior
and outer angle of each egg in reference to the mother and in the
middle line in reference to the embryo. The anterior end flattens
itself very slightly dorso-ventrally and becomes broader than the
posterior. Two pairs of prominences appear anteriorly ; the ante-
rior of which are very small and weakly developed, frequently noth-
ing more than the smallest angles on this end of the embryo. The
more posterior and larger are more sharply pronounced. They
project backwards on either side from the anterior flattened area
and make the flattening effect much more pronounced. Both pairs
of these prominences lie in a plane perpendicular to one cutting the

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118 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

gastrula month and making the embryo bilaterally symmetrical.
The first pair of these prominences become the first or sensory
antennae, while the second will be the chief locomotor organ of the
adult, the second or swimming antennae.

Up to this time the position of the embryos has been practically
the same and only a very slight change has occurred from that
first noticed in the segmenting eggs. The embryos have elongated,
and that is all. As the appendages appear the change in shape of
the embryos and of the enlarging brood-chamber causes a shifting
in position of the embryo with reference to the mother.

Upon looking at the cross-section of an adult with a brood-
ehaniber as previously described it is seen to be heart-shaped with
the apex upwards at the dorsal wall and the intestine occupying
the depression in the blunt end of the heart.

Before any change in the outward shape of the embryos occurs
they lie for the most part in the concavity on either side of the
intestine with the posterior ends of the ones in front to the outside
and somewhat beneath the anterior ends of the embryos directly
behind. When the appendages push out the flattening at the
anterior end occurs, and a resistance is encountered on either side
by growing against the outer walls of the brood-chamber externally
and against the wall of the intestine internally. The outer an-
tennae strike the wall at an angle so that an increase in width
causes the edge of the embryo to rise and revolve slightly on its
other antenna, as an axis, in such a way as to cause the region
once occupied by the gastrula mouth to look dorsally and outward
instead of outward with a slight ventral tendency. An exception
to this sometimes occurs in the case of the most anterior embryos
which do not have the other factor ; i. e. the posterior end of
another embryo under the outer side of its anterior end and chang-
ing its position. These embryos in some cases may assume such a
position that the angle formed by the antenna and the outer wall
of the animal is in the opposite direction from the rest. This
causes a rotation in the other direction so the position once occu-
j)ied by the gastrula mouth is downward and in, instead of upward
and out, as in the rest of the embryos. This opposite rotation
seems to affect only tlie most anterior embryo and even in thia
case it is the exception and not the rule. The normal or usual
position assumed by these anterior embryos is not exactly like that
taken by the rest in the brood-chamber but one in which the ven-

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SUDLER: DEVELOPMENT OF PENILIA. 119

tral surface is farther away from the mid-line and more toward the
side. Thus it is possible to distinguish almost exactly the com-
parative importance of the two mechanical factors most concerned
in the revolution.

The embryos now increase in length so that the posterior end of
each embryo lies still further back and to the outside of the anterior
end of the embryo just behind it. The height of the brood-chamber
increases, and still further growth and flattening of the embryo
caused by the enlargement of the second antennae tend to raise
the anterior of each embryo above the intestine and allow the
smaller rounded end to drop so that it now lies to the outside and
beneath the anterior end of the embryo immediately behind it.
This constantly increasing factor adds to the rotation until it may
cause the side which was originally next to the outer wall to lie on
top of the intestine. With the increase in size in all directions the
embryos become very much crowded in comparison with their condi-
tion at first. The brood-chamber also enlarges, and the floor is
raised until it is on a level with the intestine. A cross-section of it
now shows it to be hemispherical in shape rather than heart-shaped.
The embryos now lie almost wholly above the intestine and have
lost to some extent the slant in the dorso ventral plane. The
crowding occasioned upon the maturity of the embryos causes many
changes of an irregular nature in their positions, so that they depart
from the positions described, especially in regard to the ventral
side being uppermost or directly dorsal. In nearly mature embryos,
however, their long axes are more nearly parallel with that of the
mother than before. The position of the embryos of Penilia is thus
seen to be a changing one in the different stages of their develop-
ment. To a certain extent, the degree to which the embryos are
shifted depends upon their age, but throughout their development
the anterior end of the embryo lies toward the corresponding end
of the mother. In older stages, the ventral side is uppermost and
looks toward the side, except as noted, where it is just the opposite,
looking downward and toward the intestine of the mother. While
the axis of the embryo is never exactly parallel to that of the
mother, in a general way it always lies in the same direction,
sufliciently so to see with perfect clearness that the egg is oriented
and that as it develops its })osition in regard to the mother is
changed by a series of mechanical factors resulting from the
growth and the enlarging brood-chamber to accommodate its
increasing contents.

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120 PROCEEDINGS: BOSTON SOCIETY NATUBAL HISTORY.

The Appendages.

The order of the appearance pf the appendages of Penilia departs
from the usual rule for Crustacea. The first or sensory antennae
do not appear first but only after the second antennae are clearly
distinguishable, and in some cases where the retardation is excep-
tional, they may not appear sharply differentiated until the second
antennae, mandibles, and maxillary region are plainly formed. A
maxillary region is differentiated before the thoracic appendages
appear and the appendages themselves do not develop until rudi-
ments of every other appendage of the entire animal are developed.
The thoracic appendages appear in the usual order, as the most
anterior is the oldest and those following show a perfect gradation
in their development. The appendages will be taken up in the
order of their position and not in reference to the time of their
appearance.

First Antennae.

The first or sensory antennae, as has been just said, vary consider-
ably as to the time of their appearancfe. They begin as angles on
either side of the rounded anterior end of the larvae. They may
appear in this way when the second antennae are just beginning to
be plainly visible or they may not appear until the manSibles and
maxillary region are marked off. This angle increases in prominence
until it gradually forms a rounded lump projecting anteriorly, its
connection with the body of the embryo being the smallest part
of its diameter. (Fig. 28.) Its relation to the entire animal is
shifted as the bending of the head and the outgrowth of the upper
lip occur. From its extreme anterior position in front of the second
antennae it changes to the ventral side of the embryo, and finally in
the older embryos and adult its position is ventral and posterior to
that occupied by the second antennae. The first antennae project
on either side from outgrowths or horn-like processes developed
from a part of the upper lip; They remain small throughout life
and in the adult consist of but a single joint with nerve-cells and
sensory fibers or setae.

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8UDLER : DEVELOPMENT OF PENILIA. 121

Thb Second Antennae.

One of the first changes noticed in the outward shape of the
embryo is a widening of the anterior third and the appearance of
two rounded prominences projecting posteriorly on either side of
this area which will form the second antennae of the adult. These
two projections grow backward rapidly and, at a stage when the
maxillary region and two thoracic segments are present, bifurcate.
Their position in regard to the entire animal shifts, approaching
more and more the anterior end of the embryo as the head bends
and in the adult coming to occupy the extreme anterior dorsal
corner of the animal. From their first appearance the second an-
tennae are the most prominent appendages. They grow rapidly
and appear first and thus foreshadow their great importance in the
adult. The swimming hairs do not begin to show until just before
the embryos are mature. The second antennae are folded back
close to the side of the embryo as long as they are in the brood-
chamber of the mother.

The Mandibles.

The mandibles are the next appendages to appear and the time
when they are first recognizable varies but little. At a stage
represented by Fig. 21, when the second antennae are well
defined as two processes projecting posteriorly, the mandibles show
as paired outgrowths just posterior and ventral to the second
antennae. These eminences do not project either posteriorly or
anteriorly but directly outward from the middle line. As the
embryo grows older they assume an upright position (project ven-
trally) and approach more and more the middle line. In the adult
they meet in the middle line. They are stout, serrated on their
distal borders, and moved by well-developed muscles. (Fig. 1.)
Except for the change in position in the direction in which the
appendages point, there is no shifting such as one sees in the case
of the two previous appendages described. Their position in the
adult is practically that occupied by them throughout the embryonic
development.

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122 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

The Maxillae.

A maxillary region is differentiated immediately after the man-
dibles are well formed, and upon this the appendages appear later.
The first pair of maxillae appear first and the second pair follow
shortly. The first are the largest and show themselves as two
small rounded eminences projecting at first away from the middle
line but closer to it than the mandibles. Their position as devel-
opment proceeds changes but little, except that the appendages of
the adult point more toward the middle line. They are small even
in the adult and consist of but a single joint bearing setae on its
terminal extremity. The second maxillae develop at first as small
thickenings which increase until the appendages are small projec-
tions, reaching their maximum development in a stage represented
by Fig. 27, when the thoracic appendages are just beginning to show
the first trace of bifurcation. These appendages disappear as rapidly
as they come, and by the time the bifurcation of the thoracic append-
ages is well marked they have become reduced to mere darker stain-
ing masses which soon cease to be recognizable as appendages. The
second maxillae are formed sooner in Penilia than in Moina, where
they appear after the thoracic appendages have already bifurcated.
They persist, however, much longer in Moina, disappearing only just
before the larvae are set free. They are much more transient in
Penilia and never so well marked.

The TnoRA( ic Api»KNi)A(iEs.

The thoracic appendages follow the usual rule and appear in a
regular sequence of order making the anterior the oldest and the
posterior the youngest. Tlie thoracic region becomes divided into
six segments in the same order. Furrows running from the ventral
side up to the dorsal si«le ai)j)ear. These cause the prominences
between them to become more i)roiiounced as development proceeds.
This condition causes the peculiar lluted appearance presented by the
embryo in side view as seen in Fig. 2(3. These i)rominences ap})roach
much closer to the middle line on the dorsal than on the ventral side.
A view of the embryo at this stage shows the prominences projecting
slightly beyond the ventral surface and squarish in outline. This

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SUDLER: DEVELOPMENT OF PENILIA. 123

portion represents the beginning of the thoracic appendages. Near
the center of these squarish limb-rudiments which represent the
basopodite a portion becomes more prominent, projecting directly
downward. This eminence is the beginning of the endopodite and
first shows itself in the most anterior thoracic appendage, although
the difference in the time of their development is not so marked as
that of the first appearance of the thoracic segmentation. The endo-
podite grows ventral wards until we have it standing nearly at right
angles with the exopodite. The free ends of these bend more
toward the mid-line as they grow older and the shell grows back
covering them. All of the thoracic appendages are bifurcated in
the adult. The third of the series of thoracic apj)endages becomes
the largest in the adult and they grade off from it smaller in either
direction. The sixth or last pair is the smallest but preserves the
structure typical of them all.

The presence of six pairs of thoracic appendages is the chief dis-
tinction of the genus Penilia, and Claus states in reference to the
presence of six thoracic appendages in a larva of Leptodora hya-
Una that we must consider this to be the original number for the
Cladocera.

The Shell.

The shell of Penilia appears as a lobed fold of ectoderm on either
side just above the maxillary region at a stage intermediate be-
tween those shown in Figs. 1, 5, 2S. Tlie integument here rises
up into a fold projecting j)osteriorly. This fold is lobed, as seen
from a dorsal view, with the apex of tlie riiiL?ular area dividing
the two shell-rudiments directed toward tlie head of the embryo.
This is shown by the most anterior embryo of Fig. 1. On either
side this fold runs well down to the vential surface. Seen from
the ventral side when young, it resenil>K*s an aj)]»endau:e to a
considerable extent, since it is about the same width and com-
posed of two layers of cells and lies in the same general position
as that assumed by a thoracic a[)pendage. These saddle-folds
grow back gradually on either side, covering the body and apj>en-
dages as we have it in the adult. The rate of growth and shapes
can be seen in Figs. 1, 5, 6, 2><, and 29.

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124 PROCEEDINGS: BOSTON SOCIETY NATURAL mSTORY.

The Nervous System.

The nervous system of Peniliit is composed of a supraejBophageal
ganglion and a series of ventral ganglia, but the supraesophageal and
its commissures contain most of the nervous . material of the adult.
As the most important part of the nervous system, it is the first to
develop. At a stage when the second antennae, are plainly seen,
represented by Fig. 20, the ectoderm is found to possess several
layers on either side of the mid-line and at the anterior extremity of
the embryo. These paired enlargements or accumulations of cells
are the first beginning of the supraesophageal ganglion. The cells
that compose them have no distinct walls, but in some sections
appear almost circular, except where pressed against one another.
The cytoplasm is liglit staining and the nuclei are very distinct.
According to Samassa, the supraesophageal ganglion of Moina arises
from paired rudiments in the same position as described for PeniUa,
but earlier in its development. Grobben found in Moina that at
first the rudiment of the supraesophageal ganglion was single, but
afterwards became paired. He found this unpaired rudiment to
appear even earlier than Samassa, and the latter says in regard to
this that Grobben was mistaken in the stage of his embryos in
which he first found it.

The ventral chain of ganglia also rise from paired thickenings of
ectoderm. These appear externally as two slight ridges on either
side of a groove, running down the mid-line on the ventral side of
the embryo just anterior to the stomodaeum to the abdominal part
of the embryo. Grobben calls this groove the " primitivfurche "
in Moina. At a stage represented by Fig. 29, the cells composing
these two folds have separated themselves into nine groups, which
can be seen in longitudinal sections. The third one from the
anterior end of the group is the smallest and least well marked.
This is the ganglion of the second maxillar segment, and probably
fuses with the one just anterior to it, as Grobben describes for
Moina. This condition corresponds almost exactly with the figure
given by Grobben for Moina, except in that case only eight ganglia
are present, which is what we should expect, Moina having one
pair less of thoracic appendages. The supraesophageal commissure
is seen in sections as a more or less definite string of cells running
up to one side of the digestive tract. In the adult a cross-section

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8UDLER: DEVELOPMENT OF PENILIA. 125

of the veDtral nerve cord reveals a section shaped somewhat like a
dumb-bell, showing that while the ganglia unite they do not entirely
lose their double character.

The Eye.

The eye of the adult Penilia is quite small in comparison to that
of other Cladocera. Its development and separation from the
brain rudiment do not take place until quite late. The rudiments
of the eye of Penilia do not arise separately or around distinct
centers, but they are plainly seen to be lobed and paired to that
extent.

The Digestive Tract.

The beginning of the mid-gut of Penilia is seen when the inwan-
dering of cells occurs at gastrulation. The rod of tissue thus formed
breaks away or very nearly so from the ectoderm, but retains its
general position in the long axis of the embryo. The rod is solid,
but is not regular in outline or cylindrical at first. See Figs. 15,
16, 17, 30, 31, and 32. Later this indefinite rod assumes definite-
ness, and at a stage shown in Fig. 26 the mid-gut shows as a clearly
defined cylindrical rod arranged radially. Later when the embryo
enlarges and begins to assume the adult form, as at a stage repre-
sented by Fig. 28, the lumen appears. The mouth and pharynx
are formed from an inpitting which occui-s just under the rapidly
elongating upper lip. The origin of the anus is also from an inpit-
ting of the ectoderm at the opposite end of the embryo. Whether
this inpitting occurs at the same spot at which gastrulation occurred
or whether it is entirely new, I am unable to state definitely, but,
judging from the position of the gastrula mouth when the embryo
is revolved, I think the anus is formed dorsal to it. The anus
forms in Penilia directly in the center of the posterior end and in
the mid-line, while the position of the gastrula month when the
embryo is revolved is in the mid-line but considerably ventral to
the center of the posterior end. Grobben finds the anus of Moina
arising at the posterior dorsal angle of the embryo. He also finds
that the mouth and anus appear earlier in Moina than in Penilia.

According to Samassa we do not have an entoderraal rod as dis-

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12(> PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

tinct in Moina as in Penilia but more or less mixed with the meso-
derm. Only later the central cells of this mes-entoderm separate
themselves from the main mass to form a cylindrical and solid rod
of entoderm which later obtains a lumen as described for Penilia.
The cells lying outside of this are the mesodeim.

The Shell-Gland.

The shell-gland of the adult Penilia is a marked departure from
that organ as seen in other members of the group. Instead of
being long, tubular, and coiled, it is almost circular in outline. It
is composed of from six to ten large cells described elsewhere.
The shell-gland has its origin in the mesoderm in Penilia. Certain
cells in the mesoderm on either side of the digestive tract just
above the second maxillary segment begin to enlarge. These cells
have larger, darker staining nuclei than the surrounding mesoderm
cells, and they soon show a spherical arrangement. These cells con-
tinue to enlarge Until they form a solid sphere almost as large
as the organ of the mature animal but with no lumen present.
Later, but before they have assumed the definite histological struc-
ture of the adult, they appear around a lumen with an opening
to the exterior through the ectoderm. Grobben also confirms the
mesodermal origin of this gland for Moina, but the different shape
and histological character of the glands of the adults make their
rudiments in the respective embryos somewhat different. Kingsley
finds the homologous gland of Craiigon to be of mesodermal
origin and that later in the devel()[)nient it acquires an ectodermal
oi)ening to the exterior. Keichenbaeh in his work upon Astacus
and Ishikawa in his work on Atyephira both find this gland of
ectodermal origin and the mesoderm plan's no part in any stage of
its development.

The Cervical Gland.

Througliout the arthropods one or more glands are formed in
the neck and head region at various stages in their development.
Different investigators have given the same structure different
names such as cervical gland, neck-organ, dorsal gland, and
endusium. In Penilia a cervical gland is present here in the

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SUDLER : DEVELOPMENT OF PENILIA. 127

embryo but disappears before the animal is born leaving no trace
of its existence in the adult. In a Pen ilia embryo at a stage rep-
resented by Fig. 28 it shows its maximum development which is
quite weak in comparison with other members of the Cladocera. At
the stage referred to, it is composed of from fifteen to twenty cells,
each of which is longer than broad and larger at the distal end
than at the surface end. The whole organ is shaped somewhat
like a truncated cone with a small depression at the smallest end
where it comes to the surface. (Fig. 33.) It is situated in the
middle line at the anterior dorsal angle of the embryo. Its posi-
tion in Penilia is more anterior than in Moina. It appears about
the same time in embryos of both animals but disappears much
sooner in Penilia than in Moina, in which it is much stronger
developed than in the former.

The Ovary.

The ovary of Penilia is of mesodermal origin. I have failed
completely to find a genital cell from which the future genital
organs originate. I have been unable also to distinguish any
particular cell corresponding to the "Grobben'sche zelle " of
Samassa. Lebedinsky and Samassa both agree with me in being
nnable to find any trace of a differentiation of any kind at such an
early stage to form the genital organs. In Penilia certain meso-
dermal cells lying to either side of the intestine become larger and
their cytoplasm stains clearer than those surrounding them. These
form a row in a position occupied by the ovary in the adult.
Samassa finds the ovaries of Moina derived from four mesodermal
cells, two on either side. These cells multiply and grow backw^aid
to form the ovary. In Penilia the origin of the ovary corresponds
more closely to the origin of the same organ in Branchipus.
According to Claus it originates here as paired rudiments lying
in rows on either side of the digestive tract.

The Heart.

The heart of the adult Penilia is an oval-shaped body composed
of about thirty semilunar 8hai)ed cells. It possesses a single pair
of ostia situated toward the posterior end. The heart arises in the

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128 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

mesoderm in a position in the embryo corresponding to that occupied
by it in the adult. Two lots of cells, one on either side of the
middle line and above the intestine, become longer and tend to meet
in a mid-dorsal position. These gradually become longer and more
curved. These ends meet in the middle line and the structure of
the heart is practically complete as seen in the adult (Figs. 1 and
34).

Conclusion.

In reviewing the general features of the embryology of Penilia
certain characteristics present themselves prominently throughout
its development. The orientation of the egg is one of these.

An embryo always arises whose anterior end corresponds to the
same end of the egg and whose long axis also agrees with that of
the egg. Whether this is brought about by inherent differentiation
of the egg protoplasm or whether by external influences is impos-
sible of demonstration. The egg of Penilia divides transversely to
the long axis of the egg into two cells, and judging from the position
of these two cells, the descendants of the anterior one must have
most to do with the formation of the anterior end of the embryo and
the posterior one most to do with the formation of the posteiior end
of the embryo. The first two cells are certainly not right and left
in that their descendants form those halves of the body of the adult,
as Roux found for the frog's egg or Watase for cephalopod eggs,
but rather as Wilson found for Nereis, where the first cleavage is
also transverse to the future animal.

Grobben oriented the egg of Moina by means of the polar body
which lies in the substance of the egg. This body marks the ani-
mal pole, and the cell containing it in the two-celled embryo by its
development gives rise to the left side of the animal. •

The segmentation of Penilia is total and remains so throughout,
in marked contrast to most of the Crustacea. Lucifer and Euphau-
sia are the known exceptions in the higher forms.

Much less yolk is present in the egg of Penilia than in most
Crustacea even in nearly related fonns, and the segmentation cavity
never at any time possesses a yolk plug as is found in Moina.

One of the facts most frequently quoted from Grobben 's article is
the very early differentiation of one cell destined to give rise to the
reproductive organs. Samassa also finds in Moina the same cell,

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" SUDLER: DEVELOPMENT OF PENILIA. 129

but he is unable to distinguish it from its fellows after it has sub-
divided into eight cells. He cannot trace it as the mother-cell of
the genital organs, and from ray work on Penilia and Lebedinsky's
on Daphnia it is evident that Grobben's results are not confirmed.

Certain facts in the development of Penilia lead me to think
that it is a highly specialized rather than a primitive cladoceran.
The second maxillae appear as late and disappear much sooner than
they do in Moina. The dorsal gland is weakly developed in com-
parison with other members of the group and lingeous a shorter time.
The entoderm is nearly distinct as such from its origin and is never
at any time intermingled with the mesoderm as it is in Moina.
McMurrich thinks the origin of the entoderm and mesoderm
together is the rule for the entire crustacean group. In the method
of forming the eggs it resembles what is found in other Cladocera,
since the egg is here apparently the survivor of four ovarian cells
which unite to give rise to but one with little yolk. All of these
facts indicate a specialized type.

Penilia also presents other characteristics considered primitive for
the Cladocera. Clans finds six pairs of thoracic appendages present
in the metanauplius hatched from the winter egg of Leptodora
hyalinay which he thinks must be considered the original or primi-
tive number for the Cladocera. As we have seen, the presence of
six pairs of thoracic appendages is the chief distinguishing feature
of Penilia. If Clauses view be the correct one, then Penilia must
have branched off from the cladoceran stem quite early but yet
late enough to have inherited the peculiar yet characteristic method
of forming its eggs each from four equal ovarian cells.

I desire to express my thanks to Dr. C. P. Sigerfoos for the
material upon which this work was done. Also my indebtedness to
Prof. W. K. Brooks who suggested the work and throughout its
progress gave advice, and to whose kind interest is due much of
whatever value my results may possess ; and I also acknowledge my
indebtedness to Dr. H. McE. Knower for his kindly criticisms.

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130 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

LITERATURE.

Brooks, W. K.
*82. Lucifer: a study in morphology. Philos. trans, roy. soc. London, vol.
173, p. 67-137, pi. 1-11.
Claus, Carl.
'73. Zur kenntniss des baues und der entwickluiig von Branchipus stagnalis
und Apus cancriformis. Abhandl. kon. gesell. wissensch. Gottingen, bd.
18, p. 93-140, taf. 1-8.
Claus, C.
'70. Zur kenntniss der organisation und des feinen baues der daphniden und
verwandter cladoceren. Zeitsch. f. wiss. zool., bd. 27, p. 302-402, taf.
26-28.
Dohrn, Anton.
'09. Untersuchungen tiber bau und entwicklung der arthropoden. 3. Die
schalendrtise und die embryonale entwicklung der daplinien. Jena,
zeitsch., bd. 6, p. 277-292, taf. 10.
Gerstaecker, Adolph.

'00-'79. Crustacea. Bronn's Thier-reich.
Grobben, Carl.
'79. Die entwickelungsgeschichte der Moina rectirostris. Zugleich ein
beitrag zur kenntniss der anatomie der phyllopoden. Arb. zool. Inst.
Wien, torn. 2, 00 pp., 7 taf.
Grobben, C.
'80. Die antennendrtise der crustaceen. Arb. zool. inst. Wien, torn. 3. 18 pp.,
1 taf.
Ishikawa, Chiyomatsu.
'85. On the development of a freshwater macrurous crustacean, Atyephira
compressa, De Haan. Quart, journ. micr. sci., vol. 26, p. 391-428, pis.
25-28.
Kingsley, J. S.
'87, '89. The development of Crangon vulgaris. Bull. Essex inst., vol. 18,
p. 99-163, pis. 1-2 ; vol. 21, p. 1-42, pis. 1-3.
Lang, Arnold.

'91. Text-book of comparative anatomy. Translated by Herbert M. and
Matilda Bernard. London.
Lebedinsky, J.
*91. Die entwicklung der Daphnia aus dem sommereie. Zool. anz., jahrg.
14, p. 149-162.
McMurrich, J. P.
'96. Embryology of the isopod Crustacea. Journ. morph., vol. 11, p. 03-
154, pis. 6-9.

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SUDLER: DEVELOPMENT OF PENILIA. 131

Reichenbach, Heinrich.
*86. Studien zur entwicklungsgeschichte des flusskrebses. Abhandl. senc-
kenb. nat. gesell. Frankfurt a. M., bd. 14, 137 pp., 14 taf.
Roux, Wilhelm.

'88. Beitrftg© zur entwickelungsmechanik des embryo. Ueber die kUnstliche
hervorbringung halber embryonen durch zerstorung einer der beiden
ersten furchungskogeln sowie Uber die nachentwickelung (postregenera-
tion) der fehlenden korperhalfte. Arch. path, anat., bd. 114, p. 113-154,
246-291, taf. 2-5.
Samassa, Paul.

'93. Die keimblfttterblldung bei den cladoceren. 1. Moina rectirostris,
Baird. Archiv f. mikr. anat., bd. 41, p. 339-3^56, taf. 20-22.
Watase, S.
*91. Studies on cephalopoda. 1. Cleavage of the ovum. Joum. morph.,
vol. 4. p. 247-302, pis. 9-12.
Weismann, August.
74. Ueber bau und lebenserscheinungen von Leptodora hyalina, Lilljeborg.
Zeitsch. f. wiM. zool., bd. 24, p. 349-418, taf. 32-38.
Weismann, A.
'76. Zur naturgeschichte der daphniden. Zeitsch. f. wiss. zool., bd. 27,
p. 61-112, taf. 6-7.
Weismann, A., und Ischikawa, C.
*89. Ueber die paracopulation im daphnidenei, sowie tiber reifung und
befruchtung desselben. Zool. jahrb. Anat., bd. 4, p. 156-196, taf. 7-13.
Wilson, E. B.
*92. The cell-lineage of Nereis. A contribution to the cytogeny of the
annelid body. Joum. morph., vol. 6, p. 361-480, pis. 13-20.
Zaddach, E. G.

*41. De Apodis cancriformis Schaeff. Anatonie et historia evolutionis.
Bonnae, 1841, 72 pp., 4 pis.

Pnnted, October, 1800.

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SuoLEB. — Peailia.

EXPLANATION OF PLATES.
Abbreviations used.

Ant.

Anterior.

Artt.l.

1st antenna.

Ant. 2.

2d. antenna.

B. C.

Blood corpuscle.

Blastopore.

Br.

Supra-esophageal ganglion.

Br. Ch.

Brood-chamber.

Supra-esophageal commissure.

Cer. GL

Cervical gland.

Ed.

Ectoderm.

End.

Endopodite.

Ent.

Entoderm.

Ey.

Eye.

Ex.

Exopodite.

Fold between the two rudiments of the nervous system.

Ht.

Heart.

Int.

Intestine.

Upper lip.

Ms.

Mesoderm.

Mx.1.

1st Maxilla.

Mz.2.

2d Maxilla.

Mx.r.

Maxillary region.

Post

Posterior.

Sh.

Shell.

Sh. GL

Shell-gland.

Th. Seg, '

Thoracic segments.

in. iSeg, ' inoracic segments.

1, 2j 5, 4, 5, and 6. 1st, 2d, 3d, 4th, 6th, and 6th thoracic segments.

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SuDLEB. — Penilia.

PLATE 1.

Fig. 1. Adult seen from the side ; brood-chamber contains three young, the

most anterior of which is rotated in the opposite direction from the

one usually seen.
Fig. 2. Adult seen from the dorsal side. The most anterior embryo on the

left side has turned its dorsal side uppermost. The remainder

have assumed the typical position.
Fig. 3. Brood-chamber of adult containing eggs in the four-cell stage. Seen

diagonally ( dorso-laterally ) .
Fig. 4. Brood-chamber of adult containing four eggs in the thirty -two cell

stage. Seen from the dorsal surface.
Fig. 6. Side view of a brood-chamber showing embryos with three thoracic

segments. The most anterior embryo has its ventral surface

turned inward and down.
Fig. 6. Brood-chamber of an adult Penilia seen from the dorsal side. It

contains four embryos in an advanced stage of development ; all

of which present their ventral surface to view.

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SrnLER. l^ENIUA.

Pi. AT>: I .

pRor. Bust. S(x*.^AT. HiHT. Vol.. *JI>.

TMt MtLIOTVPC PRINTIMG CO. BOSTON.

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SuDLEB. — Penllia.

PLATE 2.

Fig. 7. Two-cell stage of the segmenting egg.

Fig. 8. Four-cell stage.

Fig. 9. Eight-cell stage.

Fig. 10. Sixteen-cell stage.

Fig. 11. Thirty-two cell stage. ^

Fig. 12. Sixty -four cell stage (section).

Fig. 13. Section of egg having about one hundred and twenty -eight cells and
showing the beginning of gastrulation.

Fig. 14. A more advanced stage of the same process.

Fig. 16. A still older stage showing the gastrula mouth at its greatest degree
of development.

Fig. 16. A section of a larva having all three germ layers differentiated.

Fig. 17. A horizontal section of a larva which has the first rudiments of the
two most anterior appendages.

Fig. 18. Ventral view of an embryo whose outward form is beginning to
change.

Fig. 19. An older stage than the preceding seen from the ventral surface and
showing the first traces of the first antennae.

Fig. 20. An older stage seen from the ventral side and showing the broaden-
ing at the anterior end and elongation of the embryo.

Fig. 21. An embryo seen from the ventral side showing the first and second
antennae and rudiments of the mandibles.

Fig. 22. An embryo with the first three pairs of appendages clearly defined
and a maxillary region marked off seen from the ventral side.

Fig. 23. A transverse section just behind the second antennae of a stage cor-
responding with Fig. 21.

Fig. 24. A transverse section of the same embryo nearer the head.

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SUDLER . — PENIUA .

JO.

' .-."<

13.

f .

Plate 2.

•h't'f

& * * ** •

0-0

v***-^

21.

I^it

pRoc. BosT. Soc.Nat.Hiht. Vol. 29.

TMt NCUOTVM PffUmMC CO.. BOSTON.

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SuDLEB. — Penilia.

PLATE 3.

Fig. 26. An eiubryo seen from the ventral side and which has five pairs of
thoracic ai>iHMHla,!<e.s in a<idition to those seen in Fig. 22.

A si lightly older stiige seen from the left side.

Ventral view of an embryo, at a stage when both the first and second
maxillae ai*e present.

Ventral view of an older larva. In this specimen the shell (Sh.) has
assumed cjuite definite proportions.

An embryo that has assumed somewhat the size and relations of
parts as seen in the adult, seen from the ventral side.

A horizontal section of an embryo at a stage a little older than repre-
sented by Fig. 21.

A horizontal section of an embryo somewhat younger than that seen
in Fig. 25.

A horizontal section to show the arrangement of entoderm, meso-
derm, and ectoderm of the thoracic region at a stage represented
by Fig. 26.

A section through the anterior dorsal angle of an embryo showing
the cervical gland at the stage of its maximum development.

A transverse section of an embryo at a st:ige corresponding to Fig. 28
to show the heail, intestine, and shell-glands.

Fig.
Fig.

26.

27.

Fig.

28.

Fig.

20.

Fig.

30.

Fig.

31.

Fig.

32.

Fig.

33.

Fig.

34.

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SUD1£R . — PeNIUA .

Plate 3.

t ' t * *

i •

* * •

• # 4

i9.

31.

r -

.32

.34-.

PRor. BosT. StK'.^AT.HisT. Vol. 'i9.

THC HCUOTVK PmNTIHO CO.. BOSTON.

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OCT^S 1899

iHt)

Prooaadlngs of the Boston Soolatj of Natural Hiatory.

Vol. 29, No. 7,
p. 133-162.

LIST OF MARINE MOLLUSCA OF COLDSPRING HARBOR, LONG

ISLAND, WITH DESCRIPTIONS OF ONE NEW GENUS

AND TWO NEW SPECIES OF NUDIBRANCHS.

By Francis Noybs Balch.

With one plate.

BOSTON:
PRINTED FOR THE SOCIETY.

0CTOB»R, 1899.

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OCT 98 1899

No. 7. — Zist of Marine MoUusca of Voldaprmg Harbor^ Long
l8la?idj with d€scriptio7i8 of one new Genus and two new Spe-
cies of Nudibranchs.

By Francis Notes Balcu, Cambbidge, Mass.

With one plate.

The following list of Mollusca — representing eight weeks*
work during August and September, 1898, and September, 1899,
at the Biological laboratory of the Brooklyn institute of arts and
sciences at Coldspring Harbor, Long Island, under the direction
of Dr. C. B. Davenport — claims no especial extent, complete-
ness, or novelty. It is put on record as a contribution toward more
exact knowledge of local distribution and variation and because the
nature of the locality gives it a certain interest. It represents a
fairly distinct facies of molluscan life — the fauna of the oyster
beds, broadly speaking. From this point of view its homogeneity
and the absence of stragglers lend it value. Probably almost every
species enumerated lives on the spot where found or in the imme-
diate vicinity. This characteristic makes it a good sample of
actual conditions of life in that interesting transition region where
the "Virginian" and "Acadian" (or "Boreal*') faunas overlap.
From this point of view it is, so far from being homogeneous,
strikingly heterogeneous.

Coldspring Harbor (Davenport, *98) is an indentation some
five miles long and one mile wide opening into Long Island Sound
some forty miles east of New York. It is shoal (for the most
part less than two fathoms), land-locked, tranquil, warm, muddy,
and subject to a constant inflow of detritus and fresh-water from
the high moraine hills surrounding it and abounding in streams.
The shores are little varied. Commonly, a narrow strip of salt
marsh leads to mud flats, to eel-grass beds, to oyster beds ; or the
steep " slides " of the carved moraine run down to a narrow beach
of loose sand and stones thinly strewn with glacial boulders which
shelves gently to the mud flats again. Genuine sand beaches are
wanting, sandy bottom is rare, though sand often underlies the silt
at the depth of a few feet. Rocky habitats other than the smooth
and inhospitable boulders are few, though piles, sea walls, and old

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134 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

X=r

=89%.

hulks furnish some crevices and foothold for certain species.
Gravel and stones cover small areas off the "points" where the gla-
cial hills make out. The rest of the bottom is mud, thickly strewn
with oyster, quahog, and "jingle" shells. The "Upper Harbor"
— almost cut off by a sand-spit — in which the specific gravity of
the water is at times lowered to l.OOG, forms a mud bed, rank
and black, covered with Ulvae, in which the brackish-water forms
abound and the transition from marine to fresh-water and to land
forms is almost imperceptible.

Professor Verrill (73) has analyzed the fauna of the New Eng-
land region according to environment, and gives a list of Mollusca
characteristic of each station. Below is given his analysis (slightly
rearranged) showing the per cent occurrence for each station at
Coldspring Harbor of his characteristic forms for each station.
1. JIarho7*s, estfiarie.s^ jwnds, or marshes. —

a. Sandy shores and bottoms =100%

b. Muddy shores and bottoms = "^^f/c

c. Oyster beds in brackish water = 91 %

d. Eel grass in brackish water = 80%

e. Submerged woodwork, etc. ; brackish water = ^"% ^
*1, Hays and sounds. —

a. Kocky shores = 74%
Sandy shores = 9.'} %
Muddy shores = 78%
Submerged woodwork, etc.
Rocky bottoms = 72%
Gravelly and shelly bottoms = 54%
Sandy bottoms = 7G%
h. Muddy bottoms =z 71 %

Ocean shores and outer waters. —

a. Rocky shores = 54%

Sandy shores = 90%

Rocky bottom = 55%

Sandy and gravelly bottom =: 59%

Muddy bottom = 3S% j J

Examination shows that from the "Harbors, estuaries, jjonds.

and marshes" 89% are juvsent at C'oldsju-ing Harbor, from the

" Bays and sounds" 74%, and from the "Ocean shores and outer

waters" only 59%. Again, from the sliore and shoal-bottom

b.
c.
d.
e.
f.

b.
c.
d.
e.

74% J

80%
68%

72%

50%

^ = 74%.

^ = 59%,

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BALCH: LIST OF MARINE MOLLUSCA.

185

(3 fathoms?) stations 80% are present, while from the deeper
waters we have only 59%. But a cross analysis according to other
characters shows a very different result, for from all the muddy
habitats we have 08%, from gravelly and shelly 68%, from
woodwork and rocky 68%, while from sandy we have 89%. This
last, however, should be thrown out entirely, since Professor Ver-
rill gives very few 8})ecies as characteristic of sandy shores and
bottoms and those few are common forms of wide distribution, all
occurring at Coldspring Harbor in mud.

The conclusion is that the molluscan fauna of Colds])ring Har-
bor, in spite of the well-marked character of the place as " muddy,"
is determined, not at all by that character, but preponderat-
ingly by the depth of water and by the factors included in the
'* enclosedness " of the place — that is, I suppose, by the tempera-
ture, the tranquility, the specific gravity, the per cent, of organic
matter, etc. It looks as though the various species would manage
somehow to be represented on almost any stretch of shore or bot-
tom provided only the water conditions be right. This conclusion
premises that shore and bottom were relatively equally explored,
which was probably the fact. My conclusion is, of course, not
inconsistent with Professor VerrilTs analysis, as his groupings
would still represent all they were intended to, viz., optima.

The Mollusca found were distributed, omitting doubtful occur-
rences or identifications as follows : —

Cephalopoda,

Decapoda, 1. . .
Gasteropoda^
Amphineura, 1.
Prosobranchiata.

Docoglossa, 1.

Ptenoglossa, 1.

Taenioglossa, 16.

(rymnoglossa, 8.

Rachiglossa, 9.

Toxoglossa, 1.

36.

Oj)isthol)ranchiata.
Tectibranchiata, 2.
Ascoglossa, 1.
Nndibranchiata.
Kladohepatica, 4.
Ilolohepatica, 2.
Pulmonata.

J3ass(>matophora, 2
PpJecypoda^

I^-otobranchiata, 3.
Filil>rancliiata, 6.
PscMulolanielli-

brauchiata, 2.
Eulamollibranchi-
ata, 23.
Total, ()3 genera, S3 species

34.

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136 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

Among these, apart from the nudibranchs, are no noveltieB or
even great rarities, but the following are not in Professor Verrill's
(78) Report on the invertebrates of Vineyard Sound : Littorina
littorea, Assiminea modesta^ Embletonia fuscctta, four species of
Turbonilla, Polycerdla duvenportiij and Corambelia depresaa^ while
the range of Aatarte undata^ Tergipea despecttcs^ Hemiaea cniciata,
and Cratena gymnota is extended west of the localities then known.

Sanderson Smith and Temple Prime (70) published a report
on the MoUuaca of Long Island which represented eleven years' col-
lecting and dredging and contained a great number of nominal
species. It gave to the north side of Long Island inside of Mon-
tauk Point the equivalents of about 111 modern species, of which
71, or 64% are in my list which also contains 17 species not in
5Smith and Prime's list. The additions are Littoriiia Httorea^
Alexia mgosotiSy four species of Turbonilla, several nudibranchs,
and Macoma sabulosa^ which last, however, is represented only by
one worn valve whose occurrence may be accidental. The gaps
in the list are perhaps more interesting than the occurrences. An
eastern port where neither Purpura lapiUus nor lAttorina irrorata
occurs is fairly well defined, geographically. Ten years ago it
might have been possible to define the spot within 60 miles by
saying it was a place where P, lapiUua was not, and X. littorea
was, found, but now the wave of the conquering European species
has spread far down toward Virginia and at Coldspring the native
competitor, Nassa obsokta, begins to yield room. The Caecidae,
Bela, Lacuna, Skenea, Missoa aculexia^ Odostoniia impressa and
0, seminuda^ Turbonilla interrupta and T. elega^iSy the Elysiidae,
Corbula, Saxicava, Siliqua, Donax, Tellina teneUa^ Pholas, all are
conspicuous for absence and another summer might possibly account
for some of them.

In the list of the Gasteropods as it stands 18 % are ** northern "
(i. e. forms characteristic of waters north of Cape Cod and usually
not extending farther south than New Jersey unless in deep water),
41 % are " southern *' (i, e. forms extending south to Hatteras or
Florida and found north of Cape Cod only in Massachusetts Bay or
those scattered colonies of southern forms which still dot the coast
up to Anticosti), while 42% are so general, so local, or so little
known in distribution as to be unassignable to either category. Of
the pelecypods 21% are '* northern." 52% are "southern,** and
27% are unassignable. Or, for the whole, 19-f-% are "northern,"

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BALCH: LIST OF MARINE MOLLUSCA. 137

45 -f- % are '* southern," and 36 -f % unassignable. I think the
similarity of the figures in gasteropods and pelecypods consid-
ered separately shows that these percentages stand for a real fact
of distribution. Two faunas overlap, the more southerly contribut-
ing rather more than twice as many species as the northerly.

The fojlowing species which do not occur at Coldspring Harbor
(so far as known) are given by Perkins ('69) as occurring at New
Haven on the opposite shore of Long Island Sound about forty
miles to the eastward : Amy da dissimilis Stimps. (= Astyris
zonalis Linsley), three specimens found among a lot of A. zonata;
Scalaria inuUistriata Say {^=,JScala m. Say), very rare; Odostoniia
deaUmta Stimps., very rare; O, impreaaa Stimps., not common;
O, seminuda Gould (= O. seminuda C. B. Adams) , very rare ;
Pleurotoma brunnea Perkins {=: J^efa plicata Adsuns) , one speci-
men thrown up by the waves ; Simnia xniiplicaUda Adams {^=.8,
uniplicata Sowb.) , one specimen probably brought up on southern
oysters; Jiiasoa aculens Stimps. (= B, aadeus Gould) , common
under stones and on algae near low water; Rissoellaf ebumea
Stimps. (= Hiasoa ehurneaf Stimps.) one beachworn specimen
taken by Dr. Perkins which Vemll (73, p. 655) remarks ** may
have been some other species .... I have seen no undoubted shells
of this species south of Cape Cod " ; Skeiiea planorbis F. & H.
(=z S, planorbis Fabr.), taken with Alexia myosotis, not com-
mon; Lacuna vincta Gould (= X. vincta Turton) , not very com-
mon ; IAttori7ia irrorata DeKay (= X. irrorata Say) , not at all
common ; Tornatella puncto- striata C. B. Adams (= Actaeon p.
C. B. Adams), dead on the beach, not common; Cylichna oryza
Stimps. (=: (Jylichnella o, Totten) , dead on the beach, very rare ;
Cyrtopleura truncata Tryon (= Pholas [Barnea) t. Say), clay and
peat at high water, not rare ; Martesia cimeiforniiH Tryon (= M.
Ciineifonnis Say) , one specimen in a pile of beach shells ; Saxicava
arctica Linn., sand near low water, not common ; Corbula contra eta
Say, sand at low water, not common ; Siliqua costata Con. (= A\
costata Say), very rare and small; Angulns polita? Tryon (=
Tdlina p. Say) , a few specimens doubtfully so referred ; KeUia
planidata Stimps., in accumulations of small shells on shore, not
common; Brachydontes hamatiis Say (= Jfytilus h. Say), abun-
dant on southern oysters, doubtful if it is naturalized.

The following species occurring at Coldspring Harbor do not
appear in Perkins's list: Chaetopleura apicidata Say, abundant;

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138 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

Littorina littorea Linn., abundant; Assiniinea niodesta Lea, locally
common ; two species of Turbonilla, rare ; Ilerrnaea cruciata A.
Ag., rare ; Cratena gyrnnota'i Couth., rare ; C. pilata Gould, com-
mon ; Terglpes despectus Johnst., rather rare ; EmhUtonia fuscata
Gould, rare ; Polycerella davenportii nobis, locally not rare ; Co-
ramhella dtpressa nobis, locally not uncommon ; Eriphyla^ lunulata
Con., rare ; ? ? Lucinafilosa Stimps., a drift shell ; Macoma sabidosa
Spengler, a drift shell ; GcinTtia inanhattensis Prime, rather rare.

Doubtless a certain allowance must be made for doubtful identi-
fications and not much importance can be attached to finds of one
or two dead specimens on the beach, but it appears that there were
resident at New Haven at least one Astyris, three Odostomias, one
Rissoa, Skenea, Lacuna, JAttorina irrorata^ Pholas, Saxicava,
Corbula, and Siliqua, which either do not live at Coldspring Harbor
or are so excessively rare or so excessively local or capricious as to
have escaped search. Any of them may yet be found there, how-
ever, and some, such as JAttorina irrorata^ probably will be. One
very abundant Chiton, a highly local Assirainea, two species of
Turbonilla, Eriphyla, and a southern variety of Gemma appear to
be legitimate inhabitants of Coldspring Harbor, but not of New
Haven. A comj)arison of the relative abundance of various spe-
cies emphasizes the difference.

In Perkins's list, omitting the non-acclimatized forms imported
on southern oysters (to which there is nothing corresponding at
Colds]>ring Harbor), of the gasteropods ^Ih — ^Jf^, are '* northern,"
48% are " southern, '* while 27-f-% are unassignable. Kather curi-
ously the pelecypods give the same figures so that the total is the
same. In the Coldspring Harbor list, omitting the nudibranchs
(which apparently were not collected by Dr. I'erkins), of the gas-
teropods 14-1-'^^, are "northern," 49— '^^ are *' southern '* while
Z^-\-f/r are unassignable; of the pelecyi)0(ls 23-1- <}{ are " nortli-
ern," 52-f-^/^ are "southern," while '24-f'^v are unassignable; or for
the total 18 -f- 9;^ are " northern " 50+ f^f, are " southern " and 81 -h ^^
are unassignable. The omission of the nudibranchs emphasizes
the southern aspect of the fauna on the face of the record, but
there may well be doubt whether the attribution of many of the
s|)ecies to northern waters only is not a mere effect of our lack of
knowledge of this group.

Doubtless figures like these give a false impression of accuracy,
since opinion may differ as to whether to class a species as " south-

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BALCH: LIST OF MARINE MOLLUSCA. 139

em '* or unassignable, but the sti'iking correspondence between the
figures for gasteropods and pelecypods and between the results for
Coldspring and New Haven (results obtained at first by independ-
ent classings of the species which were only afterwards compared)
shows, I repeat, that they represent real facts. The comparison
between Coldspring and New Haven at least is just.

The upshot is, as before, that two pretty distinct faunas in this
region overlap, the more southern one contributing a quota rather
more than twice that of the more northern one ; and further, the
increase in the preponderance of southern forms can be detected
in a range of forty miles.

In the list which follows the nomenclature adopted is that of
Dall ('86, '89, '89) wherever possible, while in one instance I have
followed Apgar ('91). The arrangement of the gasteropods is
that of Fischer ('87), excepting the nudibranchs where I have
followed Bergh ('92), and that of the pelecypods is that of Pel-
seneer ('94, '97). The only synonymy attempted in most cases is
Verrill's names in his report ('78) and Smith and Prime's names in
their report ('70), given for the sake of convenience in comparison.

I have adopted the scale of "very abundant" "abundant"
" very common " *' common " '* tolerably common " " rather uncom-
mon " *' uncommon " " rare " and " very rare " as the best available
way of describing the present condition of the mollusean population.
Too vague to serve as an absolute measure it is yet to be hoped
that it wilf prove accurate enough in relative terms to enable some
future student to detennine what changes in distribution and
adjustment of equilibrium a given number of years may have made
in the so-called '* permanent residents" of a given locality. If it
could be supplemented by a series of quantitative determinations
of the actually prevailing " mode " in a series of the local forms it
might prove a useful reference-point for future comparison.

A capital N or 8 following the synonymy will indicate that the
corresponding species was counted as " Northern " or "Southern."

CEPHALOPODA.

LOLIGIXJDAE.

Ijoligo pealli Les. Smith and Prime ('70), p. 405. Verrill
(73), p. 685. N.
One was taken in August, 1899, and specimens from the harbor

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140 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

are preserred at the State fish commission, which has a station
here. This species certainly occurs occasionally, if rarely.

OASTEROPODA.

AMPHINEURA.

POLYPLACOPHORA.

Chaetopleura apiculata (Say). Verrill (73), p. 661. Chiton
a. Smith and Prime (70), p. 892. S.

Abundant and unusually large on, and especially in, old shells,
etc., 1-3 fath.

PROSOBRANCHIATA.

DOCOGLOSSA.

Acmaea testudinalis (Muller;. Verrill (73), p. 661. Tectura L
Smith and Prime (70), p. 392. N.

One shell, empty but fresh. Doubtless occurs very rarely on
the more exposed points where beach boulders offer some poor
foothold.

Ptenoglossa.

Scala lineata (Say). Scalaria L Smith and Prime (70),
p. 394. Verrill (73), p. 660. S.

One fine specimen, empty but fresh, buried in clean, sandy mud
at low water.

Taenioglossa.

Naticidae.

^everita duplicata (Say). Verrill (73), p. 646. Natica d.
Smith and Prime (70), p. 396. S.

Common on the firmer bottoms at and below low water.

Lunatia heros (Say). Verrill (73), p. 646. Natica h. Smith
and Prime (70) , p. 395.

Common but less so than the preceding, with which it would
seem to compete.

The var. triseriata (Say) Verrill (73), p. 646. Natica t Smith

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BALCH: LIST OF MARINE MOLLUSCA. 141

and Prime (70), p. 395, is abundant in deeper water where it out
numbers either of the preceding.

Capulidae.

Crepidida convexa Say. Smith and Prime (70), p. 392. Ver-
rill (73), p. 650. S.

Common, but less so than the two succeeding species, and appar-
ently seeks deeper water. £gg8 still in the two-cell stage up to
August 20.

Crepidvla fomicata (Lam.). Smith and Prime (70), p. 392.
Verrill (73), p. 649. S.

Abundant at and below low water on Limulus, Fulgur, etc.
Eggs abundant in July and some still in 2-cell stage on August 23.

Crepidula plana Say. Verrill (73), p. 650. C. ungxdformis
Lam. Smith and Prime (70), p. 392. S.

Abundant on and in other shells, especially Fulgur car tea and F,
canaliculata,

LiTTORINIDAE.

Littoritia littorea (Linn^). N.

Still a recent arrival (having reached New Haven only in 1880),
and does not appear as yet seriously to threaten Xassa obsoleta, the
native competitor for the mud flats. Far less abundant than X,
paUiata or X. rudis^ from which it differs considerably in station,
it is yet common on the edges of marsh and mud flat, and every-
where scattered sparsely among the A\ obaoleta. It is not the
clean dark shell of Massachusetts Bay, but dingy gray and green
with vegetable growths like iV. obsoleta^ and appears to average
larger than north of Cape Cod ; the body-whorl seems niore inflated,
the suture less well marked. Bumpus's ('98) studies on the ven-
tricosity of this species seem to confirm such a tendency in southern
localities.

Littorina paUiata (Say). Verrill (73), p. 652. X. littoralis
var. jt>eco/iica Smith. Smith and Prime (70), p. 393. N.

Abundant 6n sea walls, piles, and wherever Fucus will grow.
As variable here as elsewhere.

lAttorina rudis (Maton). Smith and Prime (70), p. 392. Verrill
(73), p. 651. N.

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142 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

Abundant with the preceding. Relatively more abundant in less
brackish water. Extremely variable here as elsewhere.

RiSSOIDAE.

lihsoa mimita (Totten). Smith and Prime (*70), p. 393.
Littorinella m. Verrill (73), p. G53. N.

Very common on Ulva and in black mud, upper harbor, in water
almost fresh.

liiasoa laevis (DeKay). Smith and Prime ('70), p. 393. Litto-
riaella L Verrill (73), p. 653. ILjdrohia't I. Ven-ill ('82), p. 523.
? Rissoa stiinpsoni Smith. Smith and Prime (70), p. 393.

This and the preceding and two succeeding species are in such
confusion as scarcely to rei)ay any attempt to straighten out the
nomenclature without further knowledge. Stimpson ('05) on the
strength of the dentition and anatomy of JL ininuta retained
the genus Littorinella in his subfamily Hydrobiinae (= Amni-
colinae of (4ill) distinguished from Rissoinae on dentition alone,
but suggested a new genus — P^crobia — which Fischer ('ST) adopts.
The shell has also been referred to Paludinella, Cingula, and
Ilydrobia. As to II, laevis the case is much worse, for not only
does it share the above changes but sometimes one appears in one
genus or family and the other in another, while in truth the specific
difference of the two is at least doubtful. As the dentition and
anatomy of 11. laevis have never been examined (so far as I know),
any attempt to <leal with the matter must be a guess. All that is
said here applies to the two succeeding species. Dall recognizes
only 7ni/ifit(t, which he assigns to Rissoa, section Cingula. Whatever
m.'iy be thought of the generic classification, the ignoring of the
specific complications seems justified. Out of the hundred or so
specimens examined from Coldspring Harbor two answered to
the description of 7?. laevis ; a few answered to the description
of JL stimpson i better than to that of Ji. mi^iuta^ while many
answered equally to either; one answered better to the description
of li, latior than to either of the others. It seems to be true
that these variations (?) are local and that connecting series are
not very complete.

? liissoa stirnj)so?ii Smith. Smith and Prime ('70), p. 393,
Verrill ('82), p. 523. "^Littorinella laevis Verrill (73), p. 653.
Cf. supra.

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BALCH: LIST OF MARINE MOLLUSC A. 148

y ? Risaoalatior Stimps. Verrili (73), p. 655.
(Jf, supra,

ASSIMINEIDAE.

Assiminea inodesta (Lqq.). Verrili ('84), p. 253. Assitninea?
Smith and Prime (70), p. 393. A. f/rat/ana Leach. Verrili ('8*2),
p. 525.

This shell, rediscovered by Verrili at Newport in 1880 and iden-
tified at the time as the English species A, grayana but afterward
separated, apparently with some misgiving, and identified with
Lea's A. modesta (Lea, '45), is one of the interesting occur-
rences. It was found in fair numbers on four particular
stones in a sea wall and, in spite of careful search, nowhere
else. These stones were covered by brackish wat^r only for a
few minutes at high tide but were always darpp, being in the
shadow of the gang plank to the Laboratory float-stage. Here the
animals with their strangely aberrant fused eye-stalks and tentacles
(?), their bright red buccal mass showing plainly through the pale
flesh, and the translucent but solid chestnut shells with their appar-
ent double suture and the sigmoid line of the intestine showing
conspicuously on the body- whorl, might be found at any time to
the number of three or four. After collecting all that could be
found at one time, on the next day about the same number would
appear on the same space of five feet by three, and nowhere else.
Placed in salt water in the Laboratory, they crept out with all
speed, and if at once put back three or four times they appeared
eventually to become benumbed. There can be no doubt that the
species is well separated from A, f/raijiuia, as the teeth (PI. 1, fig. 11)
show well-marked difference, l)ut the identification with Lea's shell
seems doubtful. Neither his figure nor his description agrees very
well with my specimens, nor does Ven-ill's figure perfectly represent
them, though his description of the shell does so. He has not
described the animal, which has the foot broadly rounded in front,
obtusely pointed behind, translucent yellowish white, bearing the
operculum on the right side ; the muzzle bilobed, slightly wrinkled,
rather darker than foot, the red buccal mass plainly visible ; tenta-
cles and eye-stalks fused, forming thick, blunt, contractile peduncles,
each bearing two conspicuous large black eye-spots, one anterior
superior median, the other lateral anterior, the latter being the larger

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144 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

by one third. The other American representatives of the genus,
A, auberiaiia d'Orb. and A. conciima C. B. Adams, do not extend
north of Tampa, Fla. It certainly is not common and appears to
be very highly local, which would explain why a shell so conspic-
uous for everything but size has been so overlooked. If, as has
been suggested (Cooke, '95), the presence of haemoglobin in the
buccal mass is correlated with special muscular activity in those
parts, this Assiminea should prove a real glutton. The animal
proves itself very active on occasion.

Triforidae.

Triforia perversa (Linn^) var. nigrocincta (Adams). Verrill
(73), p. 648. Cerithium n. Smith and Prime (70), p. 394. S.

Uncommon. Under stones between tides. Locally on oyster
beds. Dall makes it a variety of the European species. The pro-
toconch of four whorls covered with a delicate yellowish red epi-
dermis is unusually regular and no reminiscences of a dexter habit
were detected.

Cerithiopsidae.

Cerithiopsia greenii (C.B.Adams). Verrill (73), p. 647. Ceri-
thium g. Smith and Prime (70), p. 394. S.

In September of 1899 one cast of the dredge on oyster beds
brought up this species alive to the number of ten, the only occasion
on which it occurred. With it were the young of St Ha terebralis
and adult specimens of Triforis in considerable numbers. This
brilliantly colored little shell is a beautiful microscopic object. The
three smooth, swollen, and irregular whorls of the protoconch, which
are set over out of the axis of the shell, are succeeded by two
whorls in which the adult beaded sculpture is gradually established,
first appearing not as connected beads but as disconnected rings.

Seila terebrans (C. B. Adams). Cerithiopsis t. Verrill (73),.
p. 648. C. terehellurn Stimps. Smith and Prime (70), p. 397. S.

Common alive on piles and Ulva, and (mostly dead) on shelly
bottom. Young of 2.2 mm. taken in the middle of September
show a protoconch of three swollen yellowish white whorls suc-
ceeded by a whorl showing distinct reminiscences of a beaded
sculpture, after which the regular corkscrew pattern comes in.

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BALCH: LIST OF MARINE MOLLUSC A. 145

Eumeta subulata Montagu. Cerithiopsis emersonii (C. B.
Adams). Smith and Prime (70), p. 397. Verrill (73), p. 648. S.
With the preceding but uncommon.

Cbbithiidae.

Bittium altematum (Say). J3. nigrum (Totten). Verrill (73),
p. 648. Cerithium sayi Menke. Smith and Prime (70), p. 394. S.

Abundant on the mud flats, on Ulva, etc. August 23d, young of
2-3 mm. and about 6 whorls were extremely abundant on black
mud and sand in a few inches of water, almost covering the surface
for some acres. 100 were easily picked from a square foot or two.
The young at this stage and until they have attained a length of
some 5 or 6 mm. show characters very different from the adult in
color (which is darker), the sculpture, and the aperture, which is
markedly different from, — and even less typically cerithioid
than, — that of the adult. This young stage, presumably the first
year's growth, is often discernible in the color and sculpture of the
spire-whorls of the adult as Gould (70) also has observed. A
parallel stage in apertures appears in some West Indian species of
Cerithium. No protoconch was observed.

Gymnoglossa.
Pyramidellidae.

Turbonilla sp.?

One specimen. Kindly identified for me by Miss Katherine J.
Bush as species designated as f in MS. of Bush and Verrill
which it is to be hoped may be published soon.

Turbonilla sp. ?

Common. Kindly identified by Miss Bush as species G."; cf.
supra,

Turbonilla sp. ?

Two specimens. Kindly identified by Miss Bush as species E ; ",
cf. aitpra,

Turbonilla sp.?

Two specimens. Kindly identified by Miss Bush as species V ;
cf, supra.

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14() PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

Symola producta (C. B. Adams) . Cheninitzia p. Smith aDd
Prime (70), p. 395. Odostoinia p. Verriil (73), p. 656.

A few specimens from Uiva in Upper Harbor. Rare.

Syniola fusca (C B. Adams). Ch€7yi7iitzia f. Smith and
Prime (70), p. 395. Odostomiaf. Verriil (78), p. 656.

Four specimens found with the preceding. Rare.

Odostomia hisuturalis (Say). Cheinnitzia h. Smith and Prime
(70), p. 395. Odostomia h. Verriil (73), p. 656. *

Tolerably common on Ulva and in black mud in shoal, brackish
water.

Odostomia trifida (Totten). Chemnitzia t. Smith and Prime
(70), p. 395. Odost07nia t. Vemll (73), p. 656.

Veiy common with the preceding. Young of this or preced-
ing species J mm. long were taken September 6 on the Ulva.

Ra(THI(1L0SSA.
MURICIDAE.

Enpleura caudata (Say). Verriil (73), p. 642. Banella c.
Smith and Prime (70), p. 397. S.

Common about the oyster beds, unfortunately. Very variable in
appearance according as the relations of lip and varix differ at
different stages of growth.

Urosaljnnx cinerea (Say). Verriil (73), p. 641. Buccimnn
plicosum Menke. Smith and Prime (70), p. 397. S.

Another common pest of the oyster beds. Egg capsules abun-
dant July 5th to August 15th.

CoLUMllELLlDAE.

Amxchis aimra (Say). Verriil (73), p. 643. Columbella a.
Smith and Prime (70) , p. 398.

var. shiiiUs (Havenel). Verriil (73), p. 644.

Verriil and Dall differ widely in their treatment of these forms.
Verriil (73) gives A. si}iillis as a distinct species, larger than A,
a /'((}•(/, more slender and more fusiform, more northern in distribu-
tion. Dall ('<^9) assigns to A. atyfra a range from Massachusetts
Bay to Georgia, while to its "dwarf form," A. si7nilfs, he assigns a
ran^e from Cape Lookout to Yucatan. Without attempting a dis-

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BALCII: LIST OF MARINE MOLLUSCA. 147

cussion, the form called by Verrill A, aiynilia is common at Cold-
8i)ring Harbor on shelly bottoms below tide marks. Among them
occm* a few specimens not connecting well with the common form
and answering to Verrill's interpretation of A, avara. Judging
from the general character of the fauna, it might be permissible to
predict that the commoner form is the more southerly.

Astijris liuiata (Say). Verrill ('78), p. 645. ('ohnnhella I,
Smith and Prime (70), p. 898. S.

Common everywhere below tides. Also on Ulva, etc. V^ery
conspicuous for its extreme variability of color-pattern and its activ-
ity.

var. (joiddlana Ag. MS. {':>i\n\\i^.)^:=i Bucrinnni irheatleyi
DeKay, is tolerably common.

Xassidae.

Xassa obsoleta (Say). Smith and Prime (70), p. 897. llyan-
assa o. Verrill (73), p. 641.

Abundant everywhere, especially on mud flats where it is dom-
inant.

Kassa vibex Say. Smith and Prime (70), p. 397. Verrill (78),
p. 640. S.

Uncommon. Capriciously distributed among X. obsolete, from
which it is not always readily distinguishable. A fairly well-
marked variety, yerha\}H=^ X. /rete/isis Perkins ('69), occurs, occa-
sionally in some numbers, on eel-grass.

Nftssa trivittatfi Say. Smith and Prime (70), p. 397. Tritia i.
Verrill (73), p. 641.

Abundant in proportion as the situation becomes too exposed for
X. obsoleta^ and in deeper water.

TURBIXKLLIDAE.

Fuhjvr carica {\Am\6) . Verrill (78), p. 640. Pi/Dfla c.
Smith and Prime (70), p. 898. S.

Common below tides, but less common than the following.

Fulfjur canaUnddtd (Linnc). Pyrt/ht r. Smith and Prime
(70), p. 398. /Si/cottjptis c. Verrill (78), p. 640. S.

Very common, but not perhaps abundant, from '2 fathoms out.

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148 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

TOXOGLOSSA.
CONIDAE.

Ma7igilia cerina (Kurtz and Stimps.). Venill ('73), p. 637.
Pleurotema c. Smith and Prime (70), p. 398. S.
One specimen from sandy mud in 7 fathoms.

OPISTHOBRANCHIATA.

TECTIBRANCniATA.

TORXATINIDAE.

Tornatina canaliculata (Say). JinUci c. Smith and Prime
(70), p. 399. Utricidtis c. Verrill (73), p. 663. S.

Not uncommon below tide marks, but usually dead. The speci-
mens fall into two well-separated groups, the first consisting of larger
shells with yellow epidermis and tapered spire, the second of much
smaller shells usually greatly eroded, the remains of the spire much
thicker and flatter, the columella tooth less conspicuous. Some of
the latter answer better to descriptions of lietusa pertenuis Mighels
than they do to those of T, canaliculata. Why the old shells should
have the epidermis usually intact while the young shells usually lack
it, is a puzzle. The same thing is noticeable in Cylichna alba
Brown, from more northern waters.

BULLIDAE.

Jlconinea solitaria (Say). Bulla 8. Smith and Prime (70), p.
399. Verrill (73), p. 662. S.

Dead shells common in dredging. Rather uncommon alive.
Occurs on marsh grass, top of sea wall, etc.

ASCOGLOSSA.
Hekmaeidae.

Jltnuaea critciata A. Agassiz MSS. Gould (70), p. 253.
Verrill (73), p. 667.

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BALCH: LIST OF MARINE MOLLUSC A. 149

Two specimens from Ceramiura and other red sea-weeds on
stony bottom off Lloyd's neck. Only once before reported, so far
as I know, viz., by Agassiz at Naushon in 1808; but Professor
Verrill informs me that he has taken it several times. Probably it
is not so mnch rare as capricious in occurrence, like so many nudi-
branchs.

NlTDIBUANTHIxVTA.

Kladoiiepatica.

cuatenidak.

? Cratena yymnota (Couth.). Hergh ('92), p. 81. Coryphella
(J. Verrill ('78), p. G()7.

These specimens, found depositing eggs on eel-grass, Aug. 26,
were unfortunately not preserved for complete identification. They
differed markedly from var. gouldii Ven-ill (Bergh ('9*2), p. 81,
:= Mo)ita(/tui g. Verrill ('78), p. 067), of which one specimen was
taken from compound tunicates. On the authority of Profes-
sor Verrill this is made a variety of C. gymnota,

(Jrntena pilata (Gould). Hergh ('92), p. 81. Montngna p.
Ven-ill ('78), p. 000. 31, vemufera Verrill ('78), p. 000. Aeolis
vermiferus Smith. Smith and Prime ('70), p. 891.

Common but of small size on hydroids from lobster-pot lines a
fathom or two below the surface. Also in dredgings of eel-grass.
On the authority of Professor Verrill Aeolis vermifents Smith is
reduced to synonymy. Specimens answering both descriptions were
taken.

Tkrgipedidak.

Tergipea despectas (Johnst.). Verrill ('78), p. 007. Bergh
('92), p. 81. N.

Rather uncommon on hydroids from lobster-pot lines.

Embletoniafuacata Gould ('70), p. 251. Bergh ('92), p. 84. N.

liather uacoraraon on hydroids from lobster-pot lines. This is
the first reported occurrence south of Cape Cod so far as I know.

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150 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

HOLOHEPATICA.

Phaxerobranchiata.
polyc'eridae.

PolycereUa davenportii sp. nov. PI. 1, figs. 2-7.

Body limaciform, slightly constricted behind the rhinophores. Length 2-
4 mm. Breadth (when creeping) about i of length. Color dirty green,
speckled with black and splashed with sulphur-yellow around and on rhino-
phores. Mantle distinct, without marginal flap, not covering the foot which
is pale yellowish and long in extension posteriorly where it tapers evenly to
a notched point. Anterior angles of foot moderately prolonged as oral palps,
anterior margin usually salient. Rhinophores long (in extension ^ length of
body), heavy, clavate, simple, contractile, not foliate, laminate, or retractile,
without sheaths. Gills three, small, rudimentary in appearance, dorsal, me-
dian, each consisting of a recurved stem bearing three posteriorly directed
branches between which is stretched, in perfect specimens, a delicate web.
The middle gill is set on the very conspicuous pulsating cardiac prominence
and shows scarcely any branching, the web being thicker than in the lateral
members. Occasionally a single rudimentar>' fourth branch anteriorly directed
appears. Dt)r8al papillae not on edge of mantle, usually one pair in front of,
one at level of, and one behind the gills. Two smaller pairs form a posterior
rosette. Papillae small (J length of contnicted rhinophores), inconspicuous.
Mouth anterior, funnel-shaped, dorsally exposed in extension, aiuied with
thin mandibular lamellae. Anus, a transveree silt, median, dorsal, just iR^hind
and under gills. Radula almost as in P. emertonii Verrill ('80-81, p. 387 ; '82. p.
548), rhachidlan tooth wanting; pleurae strongly hooked with accessory point.s,
large ; uncinl two, sickle-.shaped. Formula 2-1-0-1-2.

This odd little sea-slug is most nearly related to P. e)nertonii
Verrill ('80-81, p. 387; Bergh, '83, pi. 9, figs. 1-0 and pi. 8,
figs. 9-19), which it resembles in general organization, color, size, and
dentition and from which it differs in dentition (slightly), in the
fewness and smallness of the papillae and the relatively much larger
rhinophores, in the fact that the webbed gills are simply and singly
phinate instead of simply but doubly (ahernately) pinnate, in the
shape of the gills and number of their branches (which is much
greater in P. emert07ii'i)^ and finally in the character of the foot which
in P. emertonii is covered by the mantle. Professor VerrilTs great

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BALCH: LIST OF MARINE MOLLUSCA. 151

kindness in showing me his unpublished sketches of P, emertonii
has allowed me to satisfy myself of the very different appearance of
the two. Were it not for the great similarity of the dentition F.
davenportii would seem entitled to generic distinction on the ground
of the very different gills and the uncovered foot. No armature of
the penis could be made out in sections, but this was perhaps owing
to poor preservation.

Specimens of P. davenportii were first taken on August 16 from-
hydroids on lobster-pot lines, and again about two weeks later
appeared in jars of stones, weeds, hydroids, etc., which had been,
dredged in about 3 fath.

CORAMBIDAK.

Corambella gen. no v. PI. 1, figs. 12-15.

Form Corainbe-like (c/. Bergh, '71 and '92, and H. Fischer, '91), but more
convex and proportionately longer. Notaeum as in Corambe, but without
the anal notch. Rhiuophores not foliate laminate or branched ; tapered,
retractile, with sheaths. Anus and gills as in Corambe, viz. : anus median,
posterior, between the foot and the gill plates, which lie posteriorly on either
side between foot and notaeum, completely hidden by foot in life, (ienital
papilla anterior, left side, hidden in life. Jaws and pharyngeal bulb without
plates or other armatui-e. RadiUa large, no rhachidian tooth ; pleurae large,
twisted, the median ends bent up and back in a heavy hook ; uncini live, stout,
claw-shaped. Formula 5-1-0-1-5.

This genus is erected to contain a form closely allied to Cor-
ambe sargasaicola Bergh and C. testudinaria H. Fischer, but
not to be included within that genus because of the lack of the anal
notch and the different dentition. The anal notch is by Bergh
made a family character, yet no doubt the present form must fall
within his Corambidae.

Corambella depressa sp. no v.

Form Doris-like, much flattened in life, broadly rounded in front and bluntly
tapering posteriorly. Length 5 mm., broadth 3 mm. Color sometimes dull
brown with gi-ay irregular reticulation, but occasionally rather bright and
conspicuous, when a translucent dull blue gi*ound-col(»r is hidistinctly, coarsely,
and irregularly blotched with dirty green or greenish brown and irregulaily
scattered with small black spots, the i)attern darker and closer toward the
center, producing a radial effect. Between the blotches in every direction run
very conspicuous opaque yellowish white lines like the borings of larvae under

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152 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY

bark, beoomiiif? coarser toward the margin and heightening the radial effect
of the whole. These lines serve to pick out smartly the rhinophore oiH-ninp*.
In forniol specimens the color becomes black, and in specimens cleared and
viewed by transmitted light a coarse reticulatirm around the edge of the
nota^mm becomes very conspicuous especially posteriorly. Notaeum l(M)8e and
ami)ly covering all parts, often up-tiu-ned at the margin in life ; dorsal aspect,
which in life is smooth, has a shiigieen-like surface in preserved specimens from
the calcareous particles of the integument. Foot rather long, bluntly tai)ereil.
anteriorly strongly cordate, posteriorly roundly pt)inted in life (in i)reser\'ed
material usually emai'ginate). Oral disc only as broad as foot, strongly con-
vex anteriorly ; oral pali)8 very thick and .short. Rhinophores long (i maxi-
mum brea^Uh of specimen), evenly tapered, covered for about half the length
by a delicate white sheath. Entirely retnu'tile together with sheath. Gills,
which are most of the time hidden in living animals, lie posteriorly in the
s])iu*e between foot and notaeum, and consist of a set of simple overlapi)ing
l)lates on each side of the cardiat^ vessel which unites them. The.se, together
with the anal papilla which lies ventral and anterior to the heart in the median
line, are only exposed in a few even of the pre.st^rved specimens. On the left
fiide, just bfliind the neck, lies the genital (»pening from which the genitalia
are usually everted in preserved material even wIumi the animals were first
.stupelied with magnesium sulphate. In some .sections through this .**exual
extrovert ilure appeared stnmgly-staining plumose structures at first sugire.st-
ing gills and possibly representing a complicated armature of the penis but
not positively identified by me. No signs of mandibular lamellae, t)r of the
horny "Balken" described from the mouth cavity of Corambe, api)ear. The
radula is de.seribed HUj)ra.

l^iifortunately this inconspicuous little nudibranch, which occurred
during August in fair numbers under stones on whicli Fucus was
growing on the edge of swift water, and in dredgings of stones and
weed from *2 fathoms, was at first taken for DorideUa obscura Ver-
rill (78, pp. 400 and 0()4) and ('Hl-'8!> p. 547) and therefore only
casually studied and sketched in life. Consequently I am uncertain
what is the exact structure of the rhinophore and sheath, but
probably, as in Corambe, the sheath is longitudinally slit on its
posterior (superior) face and the rhinophore itself is, for the pro-
jecting portion, a sheet with the edges rolled inward posteriorly,
as tlie comparison of my sections with Fischer's (*91) very com-
plete account of the anatomy of Corambe testiid'marin makes it
clear thai almost all the structures bear some general resemblance
to that species. As the animal, in which periods of considerable
activity ap[)ear to alternate with periods of obstinate passivity,
creeps on the under surface of the water the rhinophores are com-
monly turned back in a graceful curve on each side. The gills are

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usually hidden, but rarely the progress of the animal is stopped, the
foot, which before extended beyond the mantle border, retracted,
and the extremely delicate and glassy gill-plates rapidly thrust in
and out first on one side then on the other, at such times closely
resembling the tips of some small beetles' wings thrust from under
the elytra. The form of the notaeum varies constantly in life, often
appearing broadest posteriorly and sometimes slightly emarginate
there but never so broad as long.

The relation of this form to VerrilTs DorUhUa ohsctint, which it
somewhat resembles superficially, is interesting but rather uncertain.
Both Fischer and 15ergh j)lace<l Doridella in the genus Coranibe on
the supposition that the anal notch had been overlooked and on the
strength of the general likeness. I5ut Verrills further descrij)tion
of the rhinophores as simple and without sheaths and of the gills as
''tufted" and situated on the right (left?) side near the end would
remove it not only from the genus Corambe but from the family of
Corambidae. As the structure of Doridella is entirely unknown,
placing it is only guess-work, l)ut if it proves to be a corambid
form, as seems likely, the distribution of this family of six species
will be curious, viz. : one s])ecies in the Sargasso Sea (Hergh, 71),
one in the Zuyder Zee (II. Fischer, '91), one in the China Sea
(Adams, '5S), one in the North Sea (Herbert, '!^<>), all these of one
genus, and two in Long Island Sound occupying the same station
and belonging to different genera.

PULMONATA.

Hassomatopiiora.

AuniClMDAE.

Alexin myosotin (Drap.). Verrill ('7v{), p. iMVl, X.

On stones at and above high-water mark. Usually near brackish
water. Sometimes associated with Assinnnea modesta. Uncom-
mon.

3fel(unpu8 lineatus Say. M. bid^ntatus Verrill (78), p. (>()2.
M, corneuH Smith and Prime (70), p. 899. S.

Common on edges of salt marsh, on grass stems, etc. The
banded variety occurs sparingly.

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154 PllOCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

PELECYPODA.

PROTOBRANCIIIATA.

NUCL'LIDAE.

N'ucula proxima Say. Smith and Prime (70), p. 3S5. Ver-
rill (73), p. 691.

Abundant everywhere on muddy or shelly bottoms below I fath.

Ledidae.

Yoldla Ihaatnla (Say). Verrill (73), p. GiS9. J^edd Umatuht
Smith and Prime (70), p. 385.

Locally common in black mud, 3-6 fath., and attaining a length
of IJ inches. The number of teeth is variable, ranging from 16
anterior and 13 posterior to 26 anterior and 24 posterior, so that the
numbers usually given on the authority of Gould (22-18) are mis-
leading. Probably, as Professor Verrill believes, they increase
with age, but it is easy to find small specimens with many and
large specimens with few teeth. In shape the specimens are all
nearly typical limattdas, the form sei)arated as I" sajjotida not
occurring.

SOLEXOMYIDAE.

^oleiiomya veliua Say. Smith and Prime (70), p. 389. Ver-
rill (73), p. 6S8.

Locally not uncommon at and below low water; occurs a foot or
so below the surface of the tine clean sand and mud in which it
lives.

FILIBP.VXCIIIATA.

AXUMIIDAE.

Anoiiiia shnph\c d'Orbigny. A. epluppium Smith and Prime
C70\ p. 384. A. (jlahva Verrill (73), p. 690. S.

Extremely abundant everywhere. In many places the dredge
comes up tilled with the dead shells to the exclusion of almost
everything else.

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BALCH: LIST OF MARINE MOLLUSCA. 155

Arcidae.

Area transversa Say. Smith and Prime (70), p. 8H4. ^Sca-
pharca t, Verrill (78), p. G91. S.

Common on shelly bottom, 2 fath. and below.

Area pexata Say. Smith and Prime (70), p. 8S5. Argina p.
Verrill (73), p. 092. S.

Occurs with the last species but not one fifth as common. Per-
kins ('09) notes the opposite condition at New Haven. Neither of
these " bloody-clams " appears to liave the deep-burrowing habit
which Lankester's explanation (78) of the presence of haemoglobin
in the blood calls for.

Mytilidae.

Mytilas edulis Liun^\ Smith and Prime (70), p. 880. Verrill
(78), p. 092. N.

Abundant everywhere, but the great "mussel-ridges" so charac-
teristic of some waters do not occur ; Gould's var. jtellucidHS is not
uncommon.

Modiola modiohts (Linn(»). Verrill (78), p. 098. Mi/tilus 7ii,
Smith and Prime (70), p. 8S0. N.

No live specimens were taken, but doubtless the species occurs
in the harbor as very fresh valves were tolerably common in the
dredge at certain spots.

Modiola jylicatula Lam. Verrill (78), p. ()98. Mytilus p.
Smith and Prime (70), p. 8S0. S.

Abundant everywhere, especially about salt marshes.

Specimens from a certain marsh near Lloyd's Harbor show curi-
ous distortion and erosion effects so constant as to resemble a real
variety; but doubtless the erosion, which is sharply confined to the
beaks which are buried in the soil, is due to the presence of humous
or other acids and probably the distortion is due to the same cause.
In the same marsh the clam shells were very curiously distorted ; cf,
infra.

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rSEUDOLAMELLIBRANCIIIATA.

OSTUEIDAK.

Ostrea virgintca Gmelin. O. virr/iniana Smith and Prime ('70),
p. 884. Verrill (78), p. ()97. O, horcalis Smith and Prime (70),
p. 8H4. O. costata Smith and Prime (70), p. 884. S.

Abundant, but largely artificially planted. Owing to the low
specific gravity of the water and the abundant food this is a famous
fattening ground and the industry is the principal one of the ])lace.
In 1898 the natural "set" was very small (as it had been for sev-
eral years), owing, as the fishermen believed, to severe thunder-
storms during the period when the " fry " were " swimming '' which
they variously estimate at from July 25 to August 7. In 1899 the
''set" was heavy. The method of cultivation here is simply to
clear the ground of '' wrack " and '* sludge " and plant " native "
oysters (mostly from Bridgeport, Conn., which has oysters in abun-
dance but no such fatten ing-ground as this) on grounds in the
outer waters sj)ecially prei)ared by ''cleaning u]> " and then spread-
ing gravel of such a size that each oyster may attach itself to a
separate bjvse histead of growing in bunches. As the young
approach marketable size they are moved farther up the harbor to
fatten.

Pkctinidae.

Pecten irradkins Lam. Smith and Prime (70), p. 884. Verrill
(78), p. 095. S.

Abundant on eel-grass beds in outer harbor. The spat covered
the eel-grass on Aug. 4th. Scallops are to some extent cultivated
like oysters here, but the industry does not attain to the importance
it does in Buzzard's Bay and on some parts of Caj)e Cod.

EULAMELLIBUANCIIIATA.

ASTAUTIDAE.

Astarte unduta Gould. Verrill (78), p. 684. A.sidcata Flem.
Smith and Prime (70), p. 887. N.

One live specimen from gravel and shelly bottom in 4 fath.
This was one of the surprises of dredging, as the species is here at

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BALCH: LIST OF MARINE MOLLUSCA. lo7

its extreme southern limit, I believe, as a shoal-water form, and 4
fath. would be above its usual habitat even in much more northern
waters. But the animal was alive and admitted of no doubt in
identification. Smith and Prime admit the species only doubtfully
on the authority of I)e Kay as " rare " and assign it a range from
Stonington north, so this is really an addition to their list. Verrill
mentions no locality west of New Haven, where Dr. Perkins found
var. liUea (=zAstarte httea Perkins).

Astarte castdjiea (Say). Smith and Prime (*7()), p. 887. Ver-
rill (73), p. OSf). N.

A few live specimens in black mud, 3-4 fath., and a few old
valves. Rare.

*n Astarte qttitdntns (4ould. Verrill {'78), j). 0^5.

It is doubtful whether the shell so identified was correctly placed,
even supposing this to be a good species. It is at all events a
marked variant from the preceding.

CllASSATKLLIDAK.

Eriphyla Innnlata (Conrad). Astarte /. Smith and Prime
(70), p. 387. Goahiia mactracea Gould, Verrill (78) , p. 085. S.

One dead specimen, fairly fresh, and several much worn valves
from hard bottom in 4 fath.

LrciNinAE.

?? J^uci/iafilosa Stimj)s. Verrill (78), p. OSii.

A very small shell, about 5 mm. high, extremely compressed
and with very conspicuous concentric lamellae, is assigned to this
species with the greatest uncertainty. It was fit first taken for the
young of Vemcs tnerccnaria^ which it much resembles, but there is
no pallial sinus, and the teeth, though so undeveloped as to be
equivocal, point to the Lucinidae. Being eroded and immature,
it is beyond positive identification and may have come from a
distance, very possibly carried by some fish.

Tellinidae.

Tellina tenera Say. Smith and Prime (70), p. 889. A?tf/f(lus
tener Verrill (73), p. 677. S.
Tolerably common in black mud, 3-4 fath.

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158 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

Macoma tenia (Say). Smith and Prime (70), p. 88&. Verrill
(78), p. 678. S.

Locally abundant in black mud below tide marks.

Macoma balthica (Linn^). M, fragilis Adams. Verrill (73),
p. 676. Tellina fusca Philippi. Smith and Prime (70), p. 389.

Dead but perfectly fresh and unusually large specimens locally
common 1-3 feet deep in black mud at low- water mark in Upper
Harbor. The animals doubtless live in numbers close by, prob-
ably deeper in the soft mud, although in more northern waters
and on harder ground it may be found alive free upon the surface.
The rather surprising combination of the heavy shell and epidermis
characteristic of muddy localities with the brilliant rose color
usually seen only in small shells from clean sand occurs.

Macoma sahulosa (Spengler). Verrill (73), p. 677. N.

One valve found by Dr. J. I. Ilamaker was so identified by him.

Semelidae.

Cumingia teUinoides (Conrad.). Smith and Prime (70), p.
388. Verrill (73), p. 679. S.

A few valves on hard bottoms in 4 fath. Rare.

Mactkidae.

Macira solidhsima Dillwyn. Smith and Prime (70), p. 388.
Verrill (73), p. 680.

Relatively not very common or very large. Lives in mud and
sand in more exposed situations.

Mactra lateralis Say. Smith and Prime (70), p. 388. Verrill
(73), p. 680. S.

Abundant in black mud below tide marks, slightly buried.

Vexeiudae.

Ventis intrcenaria Linn<!!\ Smith and Prime (70), p. 3S8. Ver-
rill (73), p. 681. S.

Very abundant >)elow tide marks. A considerable clam-fishery
exists.

Cytherea conveo'a Say. Smith and Prime (70), p. 388. Cal-
lista c, Verrill (73), p. 681. N.

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BALCH: LIST OF MARINE MOLLUSCA. 159

Rather uncommon on soft bottom, 3-4 fath.

Gemma purpurea H. C. Lea. Venus gemma Totten. Smith
and Prime ('70), p. 388. Tottenia gernma Verrill ('73), p. 682.

Locally abundant, lying on the surface of black mud and fine
gravel on the flats.

var. 7nanhattensis Prime. Venus m. Smith and Prime ('70),
p. 38S. Tottenia m. Verrill (73), p. 082. S.

A few specimens so identified occurred capriciously among the
G. ptfrpurea. Rare.

Petiuoolidak.

Petrlcohi pholadiformis Lam. Smith and Prime ('70), p. 390.
Verrill (73), p. 080. S.

Common, boring in the salt marsh among roots of grass, etc., at
top of beach.

Cardiidae.

Cardium pinnidatam Conrad. Smith and Prime ('70), p. 387.
Verrill ('73), p. 083. N.

One live and several dead specimens on hard bottom in 6 fath^
Rare.

Liocardhim 7nortoni (Conrad). (Jardium m. Smith and Prime
(70), p. 387. JLaevlcardium m. Verrill (73), p. 083. S.

Abundant in soft mud at and below low water.

Myidae.

Mga arenaria Linn6. Smith and Prime (70), p. 390. Verrill
(73), p. 072.

Abundant but not large. In the marsh before mentioned (see
under Jfodlola pllcatula) occurred a distorted variety, heavy, trun-
cated, and gaping, which resembled the circurapolar Jfga truncata
Linn, almost exactly, even to the tough and persistent epidermis. I
think it would scarcely be distinguished from specimens of M. trim'
cata lacking the epidermal tube. Perhaps adverse circumstances in
both cases have produced parallel results. Doubtless the thick
epidermis is a protection from the acids of the marsh.

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SOLENIDAE.

J^/isis americtma (Gould). Ensdtella a. Verrill (73), p. 674.
^Solen e?isis Linn^. Smith and Prime (70), p. 389.

Tolerably common in sand or clean mud at and below extreme
low water.

Pandouidae.

Cli(liop/i07'a (/ouldiana DM, ('. trilineata Verrill (73), p. 673.
PiUiilora t. Smith and Prime (70), p. 390.
Locally common in black mud, 3-() fath., with YohVut luiKitnld,

Lyoxsidae.

Lyonsiit hiiaUmt (Conrad). Smith and Prime (70j, p. 390.
Verrill (73), p. 672.

Rather uncommon. Locally on hard bottom in 3-6 fath.

Tekedidae.

Ttredo navalis Linn^. Verrill (73), p. 6(59. S.

Not uncommon. The submerged timber is honey-combed by
ship worms, but in only a few instances was the species determined.
Of these all but one were of this species.

Xylotnjafmbriata^Q^YGy^, Verrill (73), p. 670. S.

Found in an old hull.

Besides the above, fourteen species of land and fresh-water shells
occurre<l in the immediate vicinity of the Laboratory; and some of
them occasionally in company with Alexia inyosotis and Melampus
llneatH.H given above as marine.

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LITERATURE.

Adains, II., and Adams, A.

'58. The genera of recent Mollusca. 3v. London, 18.38.
Apgar^ A. C.

'91. Mollusks of the Atlantic coa«t of the United States south to Cai>e Hat-
teras. Journ. N. J. nat. hist, soc., vol. 2^ p. 76-162, pi. 2-4.
Bergh, R.

'71. Beitrftge zur kenntniss der mollusken des Sargassomeeres. Verhandl.
zool.-bot. gesell. Wien., bd. 21, Abh., p. 1278-1308, taf. 11-13.
Bergh, R.

'83. Beitrttge zu einer monographie der polyceraden, 3. Verhandl. zool.-
bot. gesell. Wien., bd. 33, Abh., p. 136-180, taf. 0-10.
Bergh. R.

'92. System der nudibranchiaten gasteropoden. Wiesbaden, 1892.
Bumpiis, H. C.

'98. The variations and mutations of the introduced Littorina. Zool. bull.,
vol. 1, p. 248-259, 14 pi.
Cooke, A. H.

'96. Molluscs. Cambridge natural history, vol. 3, p. 1-459.
Dall, W. II.
'86. Report on the Mollusca [of the '•Blake"], part 1. Brachiopoda and
Pelecypoda. Bull. mus. comp. zool., vol. 12, p. 171-318, 9 pi.
Dall, W. H.

'89*. Report on the Mollusca [of the "Blake"], part 2. Gastropoda and
Scaphop(Mla. Bull. mus. comp. zool., vol. 18, 492 pp., pi. 10-40.
Dall, W. H.
'89*>. A preliminary catalogue of the shell-bearing marine mollusks and
brachiopods of the south-eastern coa.st of the United States, with illustra-
tions of many of the species. Bull. 37, U. S. nat. mus., 221 pp., 74 pi.
Davenport, C. B.

'98. The fauna and flora about Coldspring Harbor, L. I. Science, n. s.,
vol. 8, p. 685-689.
Fi.scher, H.
'91. Recherches anatomitiues sur un molluscjue nudibranche appartenant
au genre Corambe. Bull, scient. France et Belgique, vol. 23, p. 358-398,
pi. 9-12.
Fischer, Paul.
'81-'87. Manuel de conchylioh)gie et de paldontologie conchyliologique.
Paris, 1881-87.
Gill, Theodore.
'71. Arrangement of the families of mollusks. Smith.sonian misc. coll., vol.
10, art. 2, 16, 49 pp.
Gould, A. A.
'70. Report on the Invertebrata of Massachusetts. 2d ed. comprising the
Mollusca, edited by W. G. Binney. Boston, 1870.

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h\'2 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

Kerbert, E.
'85. [Corambe.] Tijdschr. nederl. dierk. vereen., ser. 2, dl. 1, p. 5-0,
cxxxvii-cxxxviii.
Lankester, E. Ray.
'73. A contribution to the knowledge of haemoglobin. Proc. roy. soc,
Lond., vol. 21, p. 70-81, pi. 1.
Lea, H. C.

'44. A description of some new species of marine shells Inhabiting the coast
of the United States. Proc. Bost. soc. nat. hist., vol. 1, p. 204-206.
Lea, H. C.
'45. Descriptions of some new species of marine shells, inhabiting the coast
of the United States. Bost. journ. nat. hist., vol. 5, p. 286-290, pi. 24.
Pelseneer, P.

'94. Introduction k I'Stude des moUusques. M^m. soc. malacol. Belg., vol.
27, p. 31-248.
Pelseneer, P.

'97. Tralt6 de zoologle, public sous la direction de Raphael Blauchard.
Fasc. XVL MoUustiues, par Paul Pelseneer. Paris, 1897.
Perkins, G. H.
•69. The moUuscan fauna of New Haven. * * * Proc. Bost. soc. nat. hist.,
vol. 13, p. 109-130, 139-163.
Prime, Tem])le. See Smith, Sanderson, and Prime, Temple.
Smith, Sanderson, and Prime, Temple.
'70. Report on tlie Mollusca of Long Island, N. Y., and of Its dependencies.
Ann. lye. nat. hist. N. Y., vol. 9, p. 377-407.
Stlmpson, Wm.
'65. Researches upon the Hydrobilnae and allied forms. Smithsonian misc.
coll., vol. 7, art. 4, 3, 69 pp.
Verrill, A. E.
'73. Report upon the Invertebrate animals of Vineyard Sound and adjacent
waters, with an account of the physical characters of the region. Rept.
U. S. comm. fish and fisheries for 1871-72, p. 295-778.
Verrill, A. E.
*80-'8l. Notice of recent atidltious to the marine Invertebrata of the north-
eastern coast of Auierica, with descriptions of new genera and species and
critical remarks on others ; part 2. Proc. U. S. nat. nuis., vol. 3, p. 356-406.
Verrill, A. E.

'81-'82. Catalogue of marine Mollusca added to the fauna of the New
England region during the past ten years. Trans. Conn. acad. arts and
sci., vol. 5. p. 447-587, pi. 42-44, 57. 58.
Verrill, A. E.
'84. Second catalogue of Mollusca recently atltled to the fauna of the New
England coast and the a4ljacent parts of the Atlantic, consisting mostly of
deei)-sea species, with notes on others previously recorded. Tran.s. Conn,
aciul. arts and sol., vol. 6, p. 139-2J)4, pi. 28-32.
Verrill, A. E.
'85. Third catalogue of Mollusca recently fwlded, etc. Trans. Conn, acad
arts and sci., vol. 6, p. 895-452, pi. 42-44.

Prilled October, IS'jO.

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Balch. — Marine Mollusca. PLA.TE 1.

Fig. 1. A part of the radula of Polycerella emertonii Verrill (from Bergli '83).

Fig. 2. Two rows of teeth from the radula of Polycerella davenport ii. oc. 3.X
oil immersion ^K.

Fig. 3. A small specimen of Polycerella davenport ii creeping on a hydroid
. stem. The animal drawn from a specimen of li mm.

Fig. 4. Polycerella davenportii, doi-sal aspect, m. ^ mouth, o. p. = oral palps,
rh. = rhinophore, d. pap. ^ dorsal pappilla, c = cardijic prominence.
FroiA a si>ecimen of 3 mm.

Fig. 5. Polycerella davenport ii, posterior aspect, a. ^ arms, f. — foot.

Fig. 6. Polycerella davenportily enlarged outline of gills and cardiac promi-
nence, g. m. ^ medial gill, g. r. =^ right gill, g. 1. = left gill, w. =^
web, r. a. - rudimentary anterior branch.

Fig. 7. Polycerella davenportii, side view. From a specimen of 4 mm.

Fig. 8. Shell of Assiminea mmlcs^ta lA*a. After camera sketch. X35.

Fig. 9. Operculum of Assiminea niodesta Lea, drawn by W. Howe, after a
camera sketch. XSo.

Fig. 10. AsHiminia vKjdesta lje&. e. = eye spots, ped. = peduncle (fiL<ed eye-
stalks and tentax'les ?). b. m. = buccal mass. The figure fails to
show the relative sizes of the eye spots. The lateral spot should be
the larger by one third.

Fig. 11. One row of teeth from the raxlula of AsHiminea modesta Lea. After a
camera sketch, oc. i X obj.7.

Fig. 12. Dor.sal aspect of Coramhella depressa. From a camera sketch of a
formol sjH'cimen of 4 mm., oc. 1 Xob. 3.

Fig. 13. Ventral aspect of Coramhella depresaa from a camera sketch of a
specinuMi cleared in cedar oil, oc. 1 X obj. 3. m. ^ mouth, o. p. =
oral i)alps, .s. e. = st\\ual extrovert, f. gl. — foot glands (?), a. pap. =-
anal papilla, g. pi. .-. gill-plates, c. = cardiiic vessel.

Fig. 14.' Ventral aspect of Vorambella depressa rest in i^ on under surface of a
slide, from a camera sketch, oc. 1 X obj. 3. rh. = rhinaphore, rh. sh.
= rhinophe sheath.

Fig. 15. One row of teeth from the radula of Coramhella depressa. After a
camera sketch, oc. 1. X oil-immersion j'^.

' NoTK. ()i)i)ortnnity for further observation since the plate was made has shown
that this tijrure is not a successful representation of the living animal in its normal
state. The notaeum is represented as too ample and much too convex, the rhinophores
(which usually turn backward in a graceful sweep) as too stout, the foot in creeping
extends behind the mantle, while the oral palps are turned slightly forward. Neither
the characteristic vermiform lines of the notaeum nor a larj;:e irregularly trilobed
blotch which often ai)pears near the middle of the fcwt is shown. The general outline
is usually either almost rectangular or broadest posteriorly and often very slightly
cniarginate there.

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Proo. Bt>»T.ScK.:NAr.HisT. Voi^/il).

H*;.-ivT'''"»"i»'i'*. ' ^'''

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' OCT 23 1899

Proceedings of the Boston Society of Natural EUstory.

ToL. 29, No. 8,

p. 163-178.

THE BLOOD VESSELS OF THE m:AllT IN CARCHARIAS, RAJA,

AND AMIA.

By G. H. Parker and Fredbrica K. Davis.

With three plates.

BOSTON:
PRINTED FOR THE SOCIETY.

October, 1899.

Digitized by VjOOQ IC

OCT 28 1899

No. 8. — The Blood Vessels of the Heart i7i Carcharias^ I^qja^ and
Amia}

By G. H. Parker axd Frederica K. Davis,
Cambridge, Mass.

With three plates,
Introductio7i.

In mammals the blood vessels of the heart are usually disposed
in the following way : A right and a left coronary artery take
their origins from the base of the aorta and are distributed, in the
main, to the ventricular and auricular walls of their respective
sides. The blood from these arteries is collected by a set of super-
ficial veins arranged in three systems : a right, a left, and a median
one. The right system is represented by the right or small coro-
nary vein ; the median by the middle cardiac vein ; the left by the
left or great coronary vein and the posterior cardiac veins. All
these veins enter the coronary sinus, which in turn enters tlie right
auricle. In addition to these superficial veins, the walls of the heart
contain the numerous but small deep-seated vessels of Thebesius
which, according to Langer ('81), open from the ventricles and
auricles into a system of fine branches that connect with the coro-
nary arteries and veins by means of capillaries. In the case of the
veins, but not of the arteries, these connections may be by vessels
larger than capillaries, as demonstrated by Pratt ('98, p. 9*J) .

The extent to which the blood vessels characteristic of the mam-
malian heart occur in the lower vertebrates has never been fully
ascertained. Presumably the most primitive conditions occur
among fishes, and we have, therefore, undertaken the study of these
vessels in three easily accessible species : Ilaja erinacea Mitchill,
Carcharias littorctlis Alitchill, and Amia calva Linn. The work on
Kaja anci Amia was done in the Zoological laboratory of Rad-
cliffe college at Cambridge ; that on Carcharias was carried on at
the laboratory of the United States fish commission at Woods Hole,
Mass., and we take this opportunity of expressing our thanks to

« Contributions from the Zoological laboratory of the Museum of comparative zoology
at Haryard college. E. L. Mark, Director. No. 101.

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164 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

the Director of the laboratory, Prof. H. C. Bumpus, for much kind
assistance in connection with this portion of the research.

Coronary Arteries,

The coronary arteries and the vessels with which they are
directly connected reach a higher degree of complexity in the elas-
mobranchs than in any other group of fishes, and consequently
questions of terminology have arisen chiefly in connection with
these forms. The system which we shall adopt is, in the main,
an expansion of that employed by T. J. Parker ('84 and '87) in his
contributions to this subject, and its components may be briefly
defined in the following way : The irregular, longitudinal artery
by which the ventral ends of some or all of the efferent branchial
arteries of a given side are brought into communication may be
called the lateral hypobranchial artery (PI. 1, fig. 1, IChni. I.).
The arteries which leave the lateral hypobranchials on their median
sides and, after more or less transverse courses, unite with one
another in the median plane may be termed the commissural arter-
ies (corns, iv-v). The longitudinal median trunk produced by the
union of the commissural arteries may be designated the median
hypobranchial artery (Ji'brn, ni.). From the posterior end of the
median hypobranchial, the coronary arteries {cor. v.) pass off to
the heart. In the skates, there are in addition postenor coronaiy
arteries (PI. 1, fig, 2, cor.p.s.). These arise from a vessel which
is a branch of the subclavian artery and which from its proximity
to the coracoid portion of the pectoral girdle may be called the
coracoid artery (cc\L). The coracoid artery, besides giving rise
to the posterior coronary and certain small branches to the neigh-
boring muscles, may anastomose with either the median or the
lateral hypobranchial artery.

The terms defined in the preceding paragraph agree in general
with those used by Parker ('87), except in the case of the hypo-
branchials. This author, to whom we are indebted for the name
of these vessels, figin*ed and described them as branches from the
subclavians. After leaving the subclavians they give rise to the
posterior coronaries, and, according to him, either they retain lateral
positions, as in the skate, where they extend anteriorly to connect
with the ventral ends of the efferent branehials of either side, or
they unite in the median plane and give rise to a single longitudi-

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PARKER AND DAVIS : HEART BLOOD VESSELS IN FISHES. 165

nal vessel following the course of the ventral aorta. That the lat-
eral and median vessels are not homologous, as is implied in
Parker's account, is seen from the fact that in Mustelus, as Parker
himself ('87, p. 697, PL 34, fig. 1) has shown, both sets of vessels
may be present. We have, therefore, given them distinguishing
names: median and lateral hypobranchials. Moreover, neither of
these vessels can be properly considered a dependency of the sub-
clavian, for the branch which leaves that artery, and which Parker
regarded as their root, may be connected with them, as Hyrtl ('58,
p. 17, Taf. 2) has shown, by only a relatively small vessel. The
union, then, is not in the nature of a continuous trunk, but an
anastomosis, and the vessel posterior to this union must be consid-
ered in the light of an independent artery. This we have called
the coracoid artery.

In Carchxirias littoralis the lateral hypobranchial artery (PI. 1^
fig, 1, iVbrn. L) is a vessel irregular in its course but always con-
nected with the efferent branchials of the second, third, fourth^
and fifth visceral arches (II-V).i It may extend to meet those
of the sixth arch (VI), but when this occurs, the prolongation is
usually on one side of the animal only, and the system as a whole
is unsym metrical.

Lateral hypobranchials, essentially similar to those in Carcharias,
occur in Zygaena malltus and in Mnstelns stellatus according to
the figures given by Hyrtl ('7*2, Taf. 3, fig. 2, and Taf. 2, fig. 2)
and in Mustelns anUircticus as figured by Parker ('87, PI. 34, fig.
1). In these three species the vessels are figured as extending
from the second to the sixth arch.

Carcharias possesses two or at most three pairs of commissural
arteries. The most anterior pair lies in the grooves between
the second and third insertions of the ooracobranchial muscles (PJ.
1, fig. 1, cc^o hrn. 2 and 3) and parallel with the efferent arteries
going to the fourth visceral arches. We have, therefore, called
these vessels the commissural arteries of the fourth arch or more
briefly the fourth commissural arteries (corns, iv). In a corre-
sponding way, fifth commissural arteries (corns, r) can be dis-
tinguished. The fourth and fifth arteries were found in all the

1 In numbering the visceral arches we liave followed the scheme laid down by Oegen-
baur (".w. p. 457^, in which the first visceral arch is represented by the upper and lower
jaws, the second by the hyf>ld arch, the third by the first branchial arch, the fourth by
the second branchial arch. etc.

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166 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

specimens we dissected, while a sixth was less constant. When
present, this last was. connected with either the anterior or the
posterior efferent arteries of the sixth arch or with both. Occasion-
ally, as shown in the figure, it failed to reach the vessels of any
arch. The variations in this respect were often unsymmetrical.

Judging from the descriptions and figures given by various
authors, commissural arteries of the fourth arch occur in all sharks.*
The only exception to this statement is the observation made by
de Blaiuville ('11, p. 117) that in tSqualus peler/rinus the coronary
arteries probably come from the efferent arteries of the posterior
arch, presumably the sixth ; but as the origin of these vessels was
not exactly determined, this may have been a mistaken surmise.
From the evidence of previous figures and descriptions, commis-
sural arteries of the fifth arch occur in Mustelus (Ilyrtl, '7*2, p. 271 ;
Parker, '87, p. 697), in Zygaena (Ilyrtl, 7*2, p. 271), and in Scyl-
lium (Hyrtl, '72, p. 267 ; Marsliall and Ilurst, '92, p. 242), where,
however, they have been called by Hyrtl, Arteria nutriens recur-
rens (branchialis). Commissural arteries of the sixth arch, such
as occasionally occur in Carcharias, have been figured only in
Zygaena by Ilyrtl ('72, Tab. 3, fig. 2).

The median hypobranchial artery (PI. 1, fig. 1, JChrn, rn.) in
Carcharias is formed on the ventral side of the ventral aorta by the
union of the right and left fourth commissural arteries. This
vessel has no anterior branch such as Hyrtl ('72, p. 271) has
described in Zygaena under the name of Arteria thyreoidea impar,
but extends entirely in a posterior direction, and, after giving off
what Hyrtl ('72, p. 269) has called the epigastric branch (e'^a.),
becomes the ventral coronary {cor. v.). A dorsal coronary artery
(PL 2, fig. 4, cor. d.) is formed on the dorsal side of the ventral
aorta by a corresponding union of the right and left fifth commis-
sural arteries, supplemented by the sixth when they are present.
The trunk thus formed lies so near the heart that it may be called
the dorsal coronary artery {cor. d.), though it might without impro-
priety be regarded as in part a median hypobranchial.

Much the same condition as that found in Carcharias has been
described for Zygaena (Hyrtl, '72, p. 271, Taf. 3, fig. 2) and Mus-
telus (Parker, '87, PI. 34, fig. 2), except that in these fishes the

iThey are found in Scylliiira according to Hyrtl ('72, p. 207) and to Marshall and
Hurst (•1»2, p. 242); in Mustelus according to Hyrtl ('72, p. 271) and Parker ('87, p. 697);
and in Squatina. Acantliias, and Zygaena accordinj^ to Hyrtl ('72, p. 268-269 and 271).

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PARKER AND DAVIS : HEART BLOOD VESSELS IN FISHES. 167

ventral vessel is formed from the fourth and fifth commissural
arteries instead of from the fourth only, and the dorsal vessel is not
definitely shown to be present. In Scyllium, however, according
to Hyrtl ('72, p. 267) a dorsal vessel is present; but it arises
from the fourth commissural arteries which also give rise to jmired
ventral vessels. In Acanthias and Squatina Hyrtl ('72, p. 269)
describes only ventral vessels; these are, however, paired and
only the right one extends to the ventricle.

In Carcharias the epigastric artery is distributed to the muscles
which surround the pericardial space. According to Ilyrtl (72,
p. 269 and 271) this artery also occurs in Acanthias and Zygaena.
It has been identified in Squatina (Hyrtl, '72, p. 269) and in Mus-
telus (Parker, '87, p. 697), in both of which it anastomoses with
what we have called the coracoid artery, thus establishing connec-
tions between the subclavian and the median hypobranchial systems.

The ventral coronary artery (PI. 1, fig. 1, cor, v.) in Carcharias
divides into three vessels, a median, a right, and a left, and is thus
distributed over the ventral surface of the ventricle. The dorsal
coronary (PI. 2, fig. 4, cor. d.) also divides into three branches,
one of which goes to the right side of the ventricle and to the
auricle, another to the left side of the ventricle, and a third by
passing around the conus to the left anastomoses with branches of
the ventral coronary.

The coronary arteries in sharks have not heretofore been described
in detail except in the case of Selache, in which, according to the
careful account given by Pavesi ('74, p. 67, 6S), the plan of distri-
bution coincides almost exactly with that found in Carcharias.

As in the cases of other sharks, Carcharias possesses no posterior
coronary arteries.

In Raja eruiacea the lateral hypobranchial arteries (PI. 1, fig. 2,
h'brn. /.) show great diversity, extending in some instances over
the branchial region from the second to the sixth arch, while in
others they are restricted to the middle portion of this region.
Any efferent branchial artery of any arch between the second and
sixth may, or may not, connect with the lateral hypobranchial ;
connection is, however, the rule with vessels near the middle of the
series and the exception with those near the anterior and posterior
extremes. So far as these connections were concerned, none of the
specimens examined by us were bilaterally symmetrical.

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1G8 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

The lateral hypobranchial artery of the skate was early described
by Monro (1785, p. 16, Tab. 1, figs. 4, 5), who stated that it was
connected with all the efferent branchial arteries. Ilyrtl ('58,
p. 16) described it in liaja clavata as coming exclusively from
the vessels of the second branchial cleft, i. e., the cleft between
the third and fourth visceral arches ; and Parker (*84, p. 61) figured
it in Jiaja ruisiUa as connected with the efferent arteries of this
cleft and the next posterior. In our experience these differences
are quite as likely to be individual variations as to be of specific
impoitauce.

The commissural arteries in liaJa erinacta are of two kinds,
dorsal and ventral, of which only the dorsal correspond to the
commissurals in Carcharias. These dorsal vessels pass through
the coracobranchial muscle, either in company with the afferent
branchial arteries of the fourth arch (PI. 1, fig. 2, coins, d, n\), in
which case they correspond to the fourth commissural arteries in
Carcharias, or in company with those of the fifth arch, thus repre-
senting the fifth commissurals. A vessel (coins, d, H,) whose root
may possibly represent the sixth commissural is usually present, but
has never been observed to be connected with the lateral hypo-
branchial system. In no skate examined by us were both the
fourth and fifth commissural arteries present, but notwithstanding
this fact the two sets of vessels were so constant in their relation to
afferent branchials that their serial homology cannot be doubted.

The ventral commissural arteries in the skate (PI. 1, fig. 2,
coms.v.) spring from the lateral hypobranchials and pass mediad
between the coracobranchial and the coracohyoid muscles, i. e., lie
ventral to the coracobranchial muscle instead of dorsal to it. They
may be entirely absent and when present are usually unilateral,
though a trace of a companion vessel may sometimes be present
(Fig. 2). They occurred in six of the twelve specimens examined
by us ; in four they were unilateral and associated with dorsal com-
missural arteries on the same side ; in a fifth case the ventral artery
was unilateral but unaccompanied by a dorsal vessel on the same
side; and in the sixth instance the arteries were bilateral and asso-
ciated with but one dorsal artery. Dorsal and ventral arteries,
when both are present on a given side, unite near the ventral
aorta (Fig 2).

Heretofore dorsal and ventral commissural arteries have not been
distinguished, but when the figures and descriptions of the earlier

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PARKER AND DAVIS ; HEART BLOOD VESSELS IN FISHES. 169

authors are compared it must be admitted that both sets of arteries
have been previously observed. Thus in Baja clavata, Hyrtl ('58,
p. 16) described and figured the commissural arteries as lying
between the coracobranchial and coracohyoid muscles ; hence they
correspond to what we have called ventral commissurals ; and the
same may be said of the commissural arteries figured in a species of
skate by Martin (*94, Fig. 7). Dorsal commissurals have been
figured by Parker ('84, p. 62, Fig. 20) in Haja nasvta and by
Monro (1785, p. 16, Tab. 1, fig. 4) in the species of skate described
by him. In Jiaja 7ia8uta they are the fifth commissurals and in the
species figured by Monro they are double and represent commissurals
of the fourth as well as of the fifth arch.

The right and left commissural arteries in Maja erinacea converge
on the root of the ventral aorta, where they may anastomose, as in
i?q/a clavata (Hyrtl, '58, p. 16), or remain unconnected. From
each commissural artery a coronary artery extends over the bulbus
to be distributed to the ventricle. The left coronary (PI. 1, fig. 2,
cor, 8.) is larger than the right (cor. dx.) and is distributed in the
main to the ventral surface of the ventricle ; the right extends to the
dorsal surface of the bulb and ventricle and to the auricle. This
plan of distribution has already been observed in other species (jRaJa
clavata, Hyrtl, '58, p. 16 ; Raja sp.?, Martin, '94, p. 25 ; Torpedo
sp.?, Hyrtl, '58, p. 3).

From the coracoid arteries in Baja erinacea posterior coronary
arteries (PI. 1, fig. 2, cor.p,8.) may extend over the ventral face of
the venous sinus to be distributed to the dorsal face of the ventricle.
These vessels may be unilateral (Fig. 2) or paired (PI. 2, fig. 5).
Similar vessels have been described and figured in Maja clavata
(Hyrtl, '58, p. 17) and Raja nasuta (Parker, '84, p. 61).

The coracoid arteries in Raja erinacea may give off branches
which anastomose with what we have supposed may represent the
sixth commissural arteries (PI. 1, fig. 2, corns, d, t'l.), a condition
similar to what has already been found in other skates by Monro
(1785, Tab. 1, fig. 4), Ilyitl ('58, Taf. 2), and Parker ('84, p. 62,
Fig. 20) .

In Amia calva no trace of lateral hypobranchials was discover-
able. The arteries which supply blood to the coronaries come from
the fourth visceral arches, and hence correspond to the fourth com-
missural arteries (PI. 2, ^g. 3, corns, iv.). These unite in the

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170 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

median line on the ventral side of the ventral aorta and give rise
to a median hypobranchial artery (h*bm,m,). This extends pos-
teriorly, giving off a large epigastric branch (e^ga.), and then
divides to give rise to a ring vessel surrounding the aorta and to
two coronary arteries, one dorsal (PI. 2, fig. 6, cor. d.) and the
other ventral (PI. 2, ^^. 3, cor, v,) . These extend posteriorly over
the bulb to be distributed eventually to the respective faces of the
ventricle. Each coronary shows evidence of division into right and
left branches.

Judging from the figures and descriptions of various authors, the
coronary arteries of most teleostoraes conform to the plan in Amia.*
Probably in all the higher fishes the vessels leading from the gills
towards the heart are the fourth pair of commissural arteries.
Stannius ('46, p. 101), however, described these vessels in the stur-
geon as coming from the third branchial arch and hence corre-
sponding to the fifth commissurals ; but this is probably a mistake,
for, according to Ilyrtl ('55, p. 11), the vessels in the sturgeon
come from the fourth visceral arches as in other higher fishes. The
cod has been described by Jourdain ('07, p. 192) as receiving its
coronary supply from the third branchial arch, while Parker ('84,
p. 117) figures it as coming from the fifth. The vessel in the cod,
however, is so small and the ventral ends of the efferent branchials
are so crowded that the exact connections are rather matters of
interpretation than observation. Since the sturgeon and the cod
seem to be the only recorded exceptions to the general rule, and
since these, as exceptions, are of doubtful value, it may well be that
in all teleostomes the vessels that leave the gills for the heart come
from the fourth visceral arches and represent fourth commissural
arteries.

Both the right and the left fourth commissural arteries are well
developed in Amia, and the same is presumably true of the stur-
geon (Ilyrtl, '55, p. 11) and of the pike (Mtlller, '41, p. 198). In
Scomber and Pelarais the right artery is said (Hyrtl, '55, p. 11)

iThe chief exception to this statement is to be found in Orthragoriscus mola, bs
de8cril>e<l by Milne-Ed wants ('58, p. 341). Accordinp to this author, O.tnola has lateral
hyi)obranchial arteries connectinp theeiferent branchials of the first, second, and third
branchial arches. It has a ventral median hypobranchial formed from the fourth pair
of commissural arteries and giving rise to a ventral coronary artery. It further has a
dorsal median hjpobranchial formed from the sixth commissural arteries and giving
rise to a dorsal coronary artery. This description corresponds so well with what is
found in some elasmobranchs, and is so unlike what is known to occur in other teleo-
stomes that we have been tempted to call its accuracy into question. Unfortunately,
we have had no material by which to test this question.

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PARKER AND DAVIS : HEART BLOOD VESSELS IN FISHES. 171

to be noticeably smaller than the left. Only the left artery is
reported present in the following fishes : Perca fluviatilia (Cuvier et
Valenciennes, '28, p. 380, PI. 7, fig. 1), Salmo (Agassiz et Vogt, '45,
p. 125; Hyrtl, '55, p. 11; Martin, '94, p. 20), and (according to
Hjrrtl, '55, p. 11) Lota vulgaris^ Lucioperca 8a?idra, Silurus giants
Astro zitigel, Labrax lupus, Zeus faber, Xiphius gladius and the
cyprenoids Aspius, Squalius, Abramis, and Leuciscus. Martin ('94,
p. 16), however, states that it is tlie right artery which is present in
Abramis and the roach (Leuciscus) as well as in the carp. In
Ceratodus it is also, according to Spencer ('93, p. 8), the right
artery that is present, the left being entirely unrepresented. It is,
therefore, probable that in many teleostoraes the commissural por-
tion of the coronary system is unilaterally developed and that in
some cases it is dextral, in others sinistral.

The St/perjicial Veins of the Heart,

These veins, often included under the general name of coronary
veins, are relatively inconspicuous as compared with the coronary
arteries and have received correspondingly less attention. In Carcha-
rias littoralis they open into the venous sinus (PI. 3, fig, 7, sn, vn.)
by two orifices, one to the right and the other to the left of the
sin u-auricular aperture {ap.). The right orifice leads into the right
vein (u;t. cor. dx.) , which passes ventrally in the coronary sulcus and
is finally distributed in the main to the dorsal wall of the bulbus,
though a few small branches also pass to the right side of the ven-
tricle (Fig. 10, vn. cor, dx.). This corresponds very closely to the
right or small coronary vein in mammals. The left orifice is the
opening for three veins, the largest of which (Fig. 7, im. cor, s,)
passes to the left and is distributed to the left and ventral aspects of
the ventricle as well as to the ventral side of the bulbus (Fig. 10,
vn, cor, s,). This corresponds closely to the left or great coronary
vein in mammals. The two remaining veins (Fig. 7, V7i. crd, m.)
are small and are restricted to the dorsal surface of the ventricle.
They correspond collectively to the cardiac veins in mammals.

In the sharks reported upon by previous investigators the super-
ficial veins always open into the venous sinus near the sinu-auricular
aperture. In Squalus (de Blainville, '11, p. 117) and Scyllium
(Rdse,' 90, p. 34) there are said to be two such openings, and this is

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172 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

probably also true of Mustelus (Parker, '87, p. 720), but in Selache
(Pavesi, 74, p. 62) and Acanthias (ROse, '90, p. 34) the veins are
described as uniting and entering the sinus by a single opening.
Kight and left systems have been identified in Mustelus (Parker,
'87, p. 720) and in Selache (Pavesi, '74, p. 68), and in the latter,
as in Carcharias, the left system is more fully developed than the
right.

In liaja erhmcea the right coronary vein is represented by two
vessels (PL 3, fig. 8, vn, cor. dx,)^ which have separate openings
into the venous sinus, one of which lies more to the right than the
other. The vein on the extreme right has two principal branches,
the first one to the dorsal surface of the cone and the second to the
ventral surface of this organ and of the ventricle (Fig. 11, vn, cor,
ilx.). This second branch passes through the right side of the
groove which separates ventricle from cone. The vein to the left
also has two principal branches, one of which extends over the
dorsal side of the cone (Fig. 8) and the other passes over the ven-
tricle to the left of the cone to be distributed finally to the ventral
face of the ventricle (Fig. 11). In passing from their ventral
areas of distribution to their dorsal openings into the venous sinus
both these vessels lie to the right of the connection between auricle
and ventricle, i. e., in what coiTesponds to the right portion of the
coronary sulcus. We have, therefore, regarded them as together
equivalent to the right coronary vein.

The left coronary vein (Figs. 8 and 11, y/i. cor, $,) is a single
main trunk from the left portion of the ventricle and enters the
venous sinus by a single opening at a considerable distance to the
left of the sinu-auricular aperture («/>.)•

The right and the left coronary veins are distributed to the
whole of the ventral face and the outer edges of the dorsal face
of the ventricle. The central portion of the dorsal face is cov-
ered with a system of veins (Fig. 8, vn. crd. m.) which enter for
the most part into a transverse trunk extending parallel to the
posterior edge of the venous sinus. From this trunk small vessels
pass across to the venous sinus into whose cavity they open. This
system of vessels corresponds very closely to the cardiac veins of
mammals.

The openings of what are presumably the right and the left coro-
nary veins in a skate were described and figured by Monro (1785,
p. 18, Tab. 3, 37). The same was shown in Raja ritbus by Tiede-

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PARKER AND DAVJS : HEART BLOOD VESSELS IN FISHES. 1 73

mann ('09, p. 9, Tab. 1, fig, 2), who also figured the openings of
six small vessels belonging to the cardiac veins.

In Amia calva the superficial veins of the heart (PI. 3, fig. 9)
open into the venous sinus by a single orifice which lies posterior
and slightly to the right of the sinu-auricular aperture (ap,) .
From this opening two veins, the right (yn. cor, dx.) and the left
(vn, cor. 8.) coronary veins, encircle the heart at the level of
the coronary sulcus and anastomose so freely on the ventral side
(Fig. 12) that a ring vessel is established. From the right coro-
nary vein (Fig. 9, v?i, cor, dx,) two branches are given off, one
anteriorly to the right side of the cone, and the other posteriorly
to the same side of the ventricle. From the left coronary vein
(vn, cor, 8,) a corresponding pair of branches is given off; that
to the cone, however, is small and more ventral (Fig. 12) in posi-
tion than its fellow of the opposite side. From the ventral anas-
tomosis of the coronary veins a single vein extends over the
median ventral surface of the ventricle (Fig. 12). This probably
represents a branch from the left coronary vein. The coronary
veins of higher fishes have generally escaped attention. Rose
('90, p. 35) mentions them as present in Pirnelodus catiis and
Tetrodoii physa^ but absent from the eel; and Martin ('94, p. 21)
states that in the salmon the right coronary vein only is present
and this opens into the auricle. That there is a unilateral condition
of the coronary veins as well as of the arteries in the higher fishes
is not impossible.

Vessels of Thebesius,

The vessels of Thebesius seem heretofore never to have been
sought for in the hearts of fishes. We have endeavored to ascertain
whether they were present in the three species which we have
studied.

On inflating the left coronary vein of a fresh heart of Car-
charxas littoralis by means of a blow^pipe, the auricle was gradually
distended with air. As the entrance of air into the auricle through
the sinu-auricular opening was carefully guarded against, such air
as found its way into the heart must have come through some
other aperture. If a heart whose auricle is distended with water be
inflated as described above, bubbles will be seen forming on the
inside of the left wall, and if the opposite wall be removed, these

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174 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

bubbles can be seen coming off freely from the inner surface
of the uninjured wall. Similar experiments on the right coronary
vein gave only negative results, and from neither the right nor the
left vein could bubbling be produced from the inner surface of the
ventricle.

As this experiment can be successfully carried out with only a
slight pressure of air, there is no reason to suppose that vessels
were ruptured, and we believe we are justified in concluding that
the left coronary veins have connections which open freely on the
inner surface of the left auricular wall. These connections can be
none other than the vessels of Thebesius. Attempts to blow from
the inside of the auricle through to the left coronary vein always
failed, doubtless because of the impediment offered by the spongy
nature of the auricular wall.

Experiments of a similar character canied out on the coronary
arteries resulted in the production of small bubbles on the inside of
the left auricular wall. This, however, was accomplished only after
very vigorous blowing and consequently demonstrates that the con-
nections between the coronary arteries and the veins of Thebesius
are much more restricted than those between the coronary veins
and these vessels, a condition already observed by Pratt ('98) in
mammals.

On inflating either the right or the left coronary vein of Hcya
erinacea with air, bubbling could also be demonstrated from the
uninjured inner surface of the auricle, but no bubbling was ever
observed from the inner surface of the ventricle.

If the single opening of the coronary veins in Amia calva be
inflated, bubbling takes place from the inner surface of the ven-
tricle as well as of the auricle. This fish was the most satisfactory
of the three species for the demonstration of the vessels of Thebe-
sius.

These experiments, in our opinion, show that the hearts of fishes
possess veins of Thebesius which open into the ventricles as well
as into the auricles and which connect more freely with the coro-
nary veins than with the coronary arteries.

Conclusions.

When the blood vessels of the heart in fishes are compared with
those in mammals, the most noteworthy feature is the striking simi-

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PARKER AND DAVIS : HEART BLOOD VESSELS IN FISHES. 175

laritj between the two sets of structures. Vessels of Thebesius
may open into the single auricle and ventricle of a fish as they open
into the paired auricles and ventricles of a mammal; and their
freer communication with the coronary veins than with the coro-
nary arteries in the fish recalls the condition found in mammals.
Morphologically the vessels of Thebesius in fishes seem to corre-
spond exactly to those in mammals.

The superficial veins of the heart in fishes are also much like
those in mammals. A right, a left, and a median system can be
distinguished, and in Carchariae the vessels representing these are
almost identical with those in the mammalian heart. In only one
important respect do the superficial veins in fishes differ from those
in mammals; in fishes these veins open into the venous sinus, in
mammals into the right auricle. When, however, it is remembered
that the venous sinus in mammals becomes incorporated in the
right aui-icle this supposed difference disappears. There is then no
reason for supposing that the veins of the mammalian heart are
not homologous with those in the heart of the fish.

The coronary arteries in fishes show less resemblance to those
in mammals than has been noticed between the other classes of
vessels, and this is particularly true of the way in which the coro-
nary arteries originate. These arteries in mammals arise from the
base of the aortic arch very near the heart ; in fishes they come
from the efferent branchial arteries at places that would corresp6nd
to positions well towards the dorsal side of the aortic arch of a
mammal. The fact that the ventral aorta of a fish carries impure
blood and the corresponding vessels in a mammal pure blood, is a
suflicient physiological reason for this difference, but it leaves the
question of the homology of these parts entirely open. Have the
coronary arteries of mammals been derived from those of fishes or
are they a new system of vessels ? The supposed absence of coro-
nary arteries from the heart of amphibians has been urged in favor
of the latter opinion, but Martin's ('94, p. 59-60) statement that
in the tadpole a system of coronary arteries essentially like that in
the fish is replaced towards the close of larval life by other coro-
nary arteries is in reality almost the first piece of positive evidence
bearing on this question. While this evidence is opposed to the
homology of the coronary arteries of higher and lower vertebrates,
it must not be forgotten that the vessels of Thebesius and the
eoronary veins of the higher and lower forms show every evidence

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176 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

of being homologous, and since the arteries constitute an integral
part of this system, it seems improbable that they alone should
have undergone such fundamental replacements as is implied by
Martin's observations on the frog. Possibly the condition in the
Amphibia is to be explained on the basis of coenogenetic changes.
However this may be, the whole question of the homology of
the coronary arteries in higher and lower vertebrates seems to us
to demand much more extensive comparative study, both anatom-
ical and embryological, than has thus far been accorded to it, before
a final answer can be reached.

Summary,

1. Vessels of Thebesius have been found to open into the auri-
cle of Carcharias, and of Raja and into the auricles and ventricles
of Amia. These vessels communicate more freely with the coro-
nary veins than with the coronary arteries. They are homologous
with the similarly named vessels in mammals.

2. The superficial veins of the heart in Amia, and particularly in
Raja and in Carcharias, are arranged in three groups corresponding
to the right coronary vein, the left coronary vein, and the middle
cardiac vein of mammals. These three groups of veins in fishes
open into the venous sinus and thus agree in this respect with the
similarly named mammalian veins which open into the right auri-
cle into which the venous sinus has been incorporated. The
above mentioned superficial veins of the fish's heart are homologous
with those in the mammaPs heart.

3. The ventral ends of the efferent branchial arteries in fishes
may be connected by a lateral hypobranchial artery. From this,
commissural arteries may pass towards the median plane ; these
may be either dorsal or ventral as in Raja, and the doi*sal ones
may be serially arranged corresponding to the fourth, the fifth, and
possibly the sixth visceral arches, as in Carcharias and Raja.
The union of the right and left commissurals gives rise to, a median
hypobratichial from which coronary arteries (anterior) are given
off. These coronary arteries differ from those in mammals chiefly
in the remoteness of their point of origin. This, however, does
not necessarily preclude honiologizing them with the coronary arte-
ries in mammals.

Posterior coronary arteries were found only in Raja and have no
homoloffues in mammals.

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PARKER AND DAVIS : HEART BLOOD VESSELS IN FISHES. 177

LITERATURE.

Aerassiz, L., et Vogt, C.

'46. Aiiatoinie des salmones. Neuchatel, 196 pp., tab. A-0.
Blainville, H. de.
'11. M6moire sur le squale p61erin. Ann. mus. d'hist. nat. torn., 18, p. 88-
135, pi. 6.
Cuvier, G., et Valenciennes, A.

'28. Hist. nat. des poissons. Tom. 1. Paris, 14 + 424 pp., pi. 1-8.
Oegenbaur, C.

'98. Vergleichende anatomic der wirbelthiere. Band 1. Leipzig, 14 + 978 pp.
Hyrtl, J.

'55. tfber die selbststeuerung des herzens. Wien, 72 pp.
Hyrtl, J.
'68. Das arterielle gef ass-system der rochen. Denks. akad. wiss. Wien.
Math. nat. cl. bd. 15, p. 1-36, taf. J -5.
Hyrtl, J.
'72. Die kopfarterien der haifische. Denks. akad. wiss. Wien Math. nat.
cl. bd. 32, p. 263-275, taf. 1-3.
Jourdain, S.
'67. Sur la structure du cceur des poissous du genre gade. Comptes rendus
acad. sci., torn. 64, p. 192-194.
Langer, L.

*81. Die foramina Thebesii iin herzen des menschen. Sitzber. akiwl. wiss.
Wien. Matli. nat. cl. bd. 82, abt. 3, p. 25-39, 1 taf.
Marshall, A. M., and Hurst, C. II.
'92. A junior courst^ of practical zoology. Tliird edition. Lon<lon, 32 + 478
pp.
Martin, H.
'94. Recherclies anatomiques et enibryologiques sur les art^res coronaires du
coeur chez les vertC'hr^s. Paris, 97 pp.
Milne-Edwards, H.
'68. Le^'ons sur la physiolo^de et ranatouiie coniparfie de rhoniuie et des
anuDaux. Tom. 3. Paris, 614 pp.
Monro, A.
1785. The structure and pliysi(»loir\' of fishes explained and compared with
those of man and other animals. Ediuburirh, 128 pp., 44 tab.
Mmier,J.

'41. Vergleichemle anatomie dor myxinoideii. Drirte fcutsetzung. Ueber
dan gefasssystem. Abhaudl. akad. wiss. Berlin, 1839, p. 175-303, tab. 1-6.
Parker, T. J.

'84. A course of instruction in zootomy. London, 24 + 397 pp.

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178 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

Parker, T. J.
*87. On the blood-vessels of Mustelus antarcticus : a contribution to the mor-
phology of the vascular system in the vertebrates. Philos. trans, royal soc.
London, vol. 177, p. 686-732, pi. 34-37.
Pavesi, P.

'74. Contribuzione alia storia naturale del genere Selache. Ann. mus. civ.
sLoria nat. Genova, vol. 6, p. 6-72, tav. 1-3.
Pratt, F. H.
'98. The nutrition of the heart through the vessels of Thebesius and the
coronary veins. Amer. journ. physiol., vol. 1, p. 80-103.
Rose, C.

*90, Beitrftge zur vergleichenden anatomie des herzens der wirbelthiere.
Morph. jahrb., bd. 16, p. 27-96, taf. 4-5.
Spencer, W. B.
'93. Contributions to our knowledge of Ceratodus. Part I. — The blood ves-
sels. Linnean soc. Nev\r South Wales. The Maclearj' memorial volume,
p. 1-34, pi. 1-6.
Stannius, II.
'46. Lehrbuch der vergleichenden anatomie der wirbelthiere. Berlin, 12 -f-
482 pp.
Tiedemann, F.

'09. Anatomie des tischherzens. Landsliut, 42 pp., tab. 1-4.

Printed., October, ISUO.

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EXPLANATION OF PLATES.

Abbreviations.

acl. Auricle.

ao. V. Ventral aorta.

ap. Auriculo-ventricular aperture.

cc*d. Coracoid artery.

cc'o brn. 1-6. Insertions of coracobrancliial muscle.

cc'o. hoi. Insertion of coracohyoid muscle.

corns, iv-vi. Commissural arteries of iv-vi visceral arches.

corns, d. iv-vi. Dorsal commissural artery of the iv-vi visceral arches.

corns. V. Ventral commi.sKural artery.

cor. d. Dorsal coronary artery.

cor. dx. Right coronary artery.

cor. p. dx. Right posterior coronary artery,

cor. p. s. Jjiift posterior coronary artery.

cor. s. Left coronary artery.

cor. V. Ventral coronary artery.

e^ga. Epigastric artery.

h'bm. I. Lateral hypobranchial artei*y.

h'brn. m. Median hypobranchial artery.

sn. vn. Venous sinus.

vn. cor. d. RJ^'ht coronary vein.

vn. cor. s. Left coronary vein.

vn. crd. m. Median cardiac vein.

vnt. Ventricle.

I-VII. Visceral arches I-VII.

The figures are drawn from actual dissection and are about natural size.

Blood vessels covered by other structures are often indicated by dotted lines.

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Parker and Davis. — Heart blood vessels in fishes.

PLATE 1.

Fig. 1. Ventral view of the blood vesselH iu the region of the heart and bran-
chial clefts of Carcharias littondis. The insertions of the coraco-
hyoid and coracobranchial muscles are indicated on the left side only.
3 and 5 third and fifth insertions of the coracobranchial muscle.

Fig. 2. Ventral view of the blood vessels in the region of the heart and bran-
chial clefts of Haja erinacea.

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Pakkbb and Davis. — Heart Blood Vessels in Fishes. Plate 1.

cc'o hoi

Proc. Bost. Sor. Nat. Hist. Vol. 29.

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Parker and Davis. — Heart blood yessels In fishes.

PLATE 2.

Fig. 3. Ventral view of the heart and the adjacent vessels in Amia calva.

Fig. 4. Dorsal view of the heart and attached vessels of Carcharias. The

auricle has been removed.
Fig. 5. Dorsal view of the heart of Raja with the venous sinus laid open and

the posterior coronary arteries shown.
Fig. 6. Dorsal view of the heart and the coronary artery in Amia. The
auricle has been removed.

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Tarkek and Davis. — Heart Blood Vessels in Fishes. Plate 2.

ao.v.
corns. IV

cot.cL.

ap.

cor.p.i. cor.p.dx.

4 6

Proo. Bost. Soc. Nat. Hist. Vol. 29.

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Pabreb and Davis.— Heart blood vessels In llsbes.

PLATE 3.

Fig. 7. Dorsal view of the heart and its superficial veins in Cafcharias. The

venous sinus is laid open.
Fig. 8. Dorsal view of the heart and its superficial veins in Raja. The venous

sinus is laid open and the auricle removed.
Fig. 9. Dorsal view of the heart and its superficial veins in Amia. The venous

sinus is laid open..
Fig. 10. Ventral view of the heart and its superficial veins in Carcharias.
Fig. 11. Ventral view of the heart and its superficial veins in Raja.
Fig. 12. Ventral view of the heart and its superficial veins in Amia.

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Pabker axd Davis. — Heart Blood Vessels in Pishes.

Plate 3.

.vn.cor.dK,

-vn.ctd.m

.vn.coT.dx.

ao.v.

vn.cof.di

A-vn.cof %.

Proc. Bost. Soc. Nat. Hist. Vol. 29.

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APR ?.0 1900

Proceedings of th» Boston Society of Natural History.
Vol. 29, No. 9,
pp. 179-184.

THE OCCURRENCE OF FOSSILS IN THE ROXBURY CONGLOMERATE.

By Henry T. Btrr ani> Rohert E. Biukk.

VViTU OXE 1*LATE.

BOSTON:
PRINTED FOR THE SOCIETY.

April, li)00.

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APR 20 1900

No. 9. — The Occurrence of Fosaila in the Roxbury Conglomerate.
By Henry T. Burr and Robert E. Burke.

The age of the Boston Basin sediments has long been a matter
for controversy. Two types of rock make up the mass of the sec-
tion, a thick series of heavy conglomerates, and a series of compact,
flinty slates. The former are known as the Roxbury conglomerates,
and are usually assumed to be of the same age throughout. The
slates are generally similar in character, but are now believed to
represent at least two widely different horizons. The conglom-
erates and much the greater portion of the slates are remarkably
barren of fossils. In 1856 Paradoxides harlani Green was reported
from the slates near Ilayward's Creek, Braintree, Rogers, *56,
pp. 27-29, 40-41. This discovery established the age of that
portion of the slates as Cambrian. As the greater part of the slates
in the Basin resemble those of Braintree, the whole series was
regarded as of the same age. Within recent years Lower Cambrian
fossils have been found in the impure limestone at Nahant, Foerste,
*89, pp. 261-263, and at Mill Cove, Weymouth, Grabau, '98, also
Burr, 1900.

In some parts of the Basin, at least, the conglomerates appear to
underlie the slates ; hence they, too, were held to be of Cambrian
age. The conglomerates are largely made up of fragments which
appear to have been derived from the complex of granitic rocks to
the south, Crosby, '89, p. 6. The granite, then, is older than the
conglomerate. On the supposition that the conglomerate is below
the slate, it is necessary to regard the granite as older than the
slate, also. But the granite, as is now known, is intrusive into the
slates of Braintree, Wadsworth, '83, p. 27 ; also Crosby, '89, p. 5,
and is, ' therefore, later than that portion of the Cambrian series.
The conglomerates, then, not only overlie the Middle Cambrian
slates, but are separated from them by a great period of igneous
action, and an interval long enough to allow the forces of erosion to
penetrate deep into the granite mass. It is, therefore, no longer
necessary to regard the conglomerates and associated slates as of
Cambrian age. In the absence of positive evidence to the contrary,
however, they are still held to be Cambrian by many observers, and
are so mapped by Walcott in his correlation papers, Walcott, '91,
p. 268, and map, p. 358.

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180 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

Of late years the opinion has been gaining ground that the con-
glomerates, at least, should be referred to the Carboniferous. This
belief is based largely upon the strong lithological resemblance
between the Boston Basin series and the Carboniferous conglom-
erates of the neighboring Norfolk County and Narragansett Basins.
The argument from analogy is not strictly valid, for it is quite pos-
sible that very similar conglomerates might be formed at widely
different horizons, particularly where the sources of supply remain
the same. In fact, as has been pointed out, our glacial material, if
worked over by the sea, would yield a deposit essentially like the
Roxbury conglomerate.

Until recently the idea of the Carboniferous age of these sedi-
ments has been practically unsupported by fossil evidence. Some
years ago Mr. J. B. Woodworth found, in the conglomerate near
Franklin Park, a fragment which he regarded as a portion of a fossil
plant, similar to forms occurring in the sandstones of the Narragan-
sett Basin. Not then appreciating the importance of his discovery,
he did not preserve the specimen. Other observers have reported
the discovery of fossils in the conglomerates and associated slates,
but no identifiable forms have ever been produced.

During the past year the writers have made a careful search over
a considerable portion of the area. Traces of possibly organic
remains have been found in the slates and in the quartzite pebbles
of the conglomerate, but none of these have had any determinative
value. Recently, however, a sandy zone near the top of the con-
glomerates has yielded fossils of a much more satisfactory nature.
These are believed to be casts and moulds of the trunks or roots of
tree-like forms. They are cylindrical m form, with circular cross-
sections. The largest has a maximum diameter of four and eight
tenths inches. They are marked by somewhat irregular transverse
wrinklings (see Plate 1), which sometimes pass entirely aiound the
form, sometimes die out, or become united. The organic matter
has entirely disappeared. The casts are composed of compact sand-
stone, which is indistinguishable from that of the rock in which they
lie. The bedding of the sandstone is obscure, but is believed to lie.
at right angles with the axes of the fossils.

The species to which these forms should be referred cannot at
present be determined. The genus, even, is highly problematical.
The only markings which are at all characteristic are the transverse
wrinklings referred to above. Quite similar wrinklings appear

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BURR AND BURKE : FOSSILS IN ROXBURY CONGLOMERATE, 181

upon several species of the genus Artisia^ particularly upon Artisia
distans, figured by Grand *Eury in Flore Carbonifer^ de la Loire.
It is quite possible that the specimens figured are related to that
form. It cannot, however, be denied that such vague markings
have little determinative value and may even be of mechanical
origin.

While the writers are disposed to believe that these forms cannot
be identified with certainty, they nevertheless feel confident that
they are true fossils. The several geologists who have passed judg-
ment upon them, have, with one exception, expressed themselves as
satisfied of their organic origin. The other view is that they are
due to mechanical action, — that they are akin to stylolites. It
is difficult to see upon what this opinion is based. The speci-
mens figured certainly bear little resemblance to the ordinary forms
of stylolites. As generally defined, stylolites are forms produced in
rocks by displacement or the development of slickensides about a
portion protected by a shell or other hard capping. Such forms
have characteristically slickensided surfaces, usually with strongly
marked longitudinal stHations, and frequently with the development
of secondary minerals. The forms under consideration show no
trace of longitudinal striations. The surfaces are not smooth after
the manner of rubbed surfaces. There is, so far as can be seen, no
development of new minerals. In short, the phenomena of slicken-
siding are altogether absent. Stylolites are of small size, seldom
exceeding four inches in length or two in diameter. The specimens
figured average about four inches in diameter, and the largest is over
a foot in length, with the total length not known. There is no
reason why stylolites should have a circular cross-section. It would
be strange indeed if, as in this case, all the specimens found in a
limited area should have this form. There seems then to be no
reason whatever for thinking that the surfaces of these specimens
were developed by differential movement. It should be added that
stylolites usually occur in limestones or in fine-grained shales, and
have never, so far as is known, l)een reported from coarse sandstones.

It may be suggested that these forms are concretionary. Cylin-
drical or rod-like concretions are not unknown, and, so far as the
form is concerned, these specimens might be of such nature. If
the forms are concretionary, the material of which they are made
up should differ, in a determinable way, from the material of the
matrix. This does not appear to be the case. In both cast and

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182 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

matrix the material is a compact quartzose sandstone, without trace
of lime and with but enough iron to give it a reddish tinge. More-
over, although similar sandstone is abundantly developed in portions
of the conglomerate area, it nowhere shows signs of concretionary
action.

No other action comes to mind as capable of producing similar
forms, unless it is assumed that they may be due to jointing. But

MAP

Showing tocd^lity
of

FOSSILS.

SCALE SOO'-l".

it seems extremely improbable that jointing should produce such
regular forms. Moreover, the surfaces lack all the characteristic
features of joints, Woodworth, '96, pp. 163-183.

All things considered, the forms are best explained as casts and
moulds, and may be fairly assumed to be of organic origin.

Granting this, it is still true that these fossils do not definitely

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BURR AND BURKE : FOSSILS IN ROXBURY CONGLOMERATE. 183

determine the age of the conglomerate. They might have come
from the Devonian or the Trias as well as from the Carboniferous.
But they surely do settle the question as between Cambrian and
Carboniferous. And this is and has been the point at issue.

The fossils were found in Forest Hills Cemetery, on the south-
em edge of the wide belt of conglomerate extending through
Brookline, Roxbury and Dorchester. The field relations show
that this conglomerate mass is, structurally, a broad, flat-topped
anticline. The fossil horizon is, therefore, at the top of the series.
The slates surround this anticline, and everywhere dip and strike
conformably with the adjacent beds of the conglomerate series. It
is a safe assumption that they overlie the conglomerates and are
conformable with them. The accompanying section shows the

FOSSIL

Conglomer^e with S*>nd»ton« bands in upper portfons. /HORIZON.

•n««NI.U«S. JAMAICA ^LAIM. POReST HtkLl

GENERAUZED SECTION ACROSS MAIN CONGLOMERATE MASS.

structural relation of the fossil horizon to the conglomerates and
the overlying slates. The evidence from these fossils seems appli-
cable to the whole of this conformable series.

It has not, as yet, been definitely proved that the other conglom-
erates of the region are of the same age as those of the central belt,
although such is generally assumed to be the case. They are much
alike lithologically, show the same degree of secondary alteration,
and have the same apparent relations with the igneous rocks. The
outcrops of slate are so scattered that it is not possible to correlate
them in a satisfactory manner. It seems probable, however, that
the greater part of the slate is closely associated with the conglom-
erate, and of approximately the same age.

It is believed, then, that the discovery of these forms serves to
establish the fact that Carboniferous sediments are present in the
Boston basin, and to render it probable that the greater part of the
area is occupied by sediments of this age.

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184 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

LITERATURE.

Burr, H. T.
1900. A new Lower CanibriaD fauna from Eastern Massachusetts. Amer.
geol., vol. 26, pp. 41-60.
Crosby, W. O.

'89. Physical history of the Boston basin. Boston, 1889, pp. 6, 6.
Foerste, A. F.
*89. Palaeontological horizon of the limestone at Nahant. Proc. Boston soc.
nat. hist., vol. 24, pp. 261-263.
Gtabau, A. W.
*98. Guide to localities illustrating the geology, marine zoology and botany
of the vicinity of Boston. Boston, 1898. (Refers to discovery by Prof.
W. O. Crosby.)
Rogers, W. B.
, *56. Notes on Paradoxides from Braintree. Proc. Bosttm soc. nat. hist.,

vol. 6, pp. 27-29, 40-44.
Wadsworth, M. E.
'83. On the relation of the Quincy granite to the Primordial argillite of
Braintree, Massachusetts. Proc. Boston soc. nat. hist, vol. 21, pp. 274-277.
Walcott, C. D.
'91. Correlation papers. — Cambrian. Bull. 81, U. S. geol. surv., p. 268,
map, p. 858.
Wood worth, J. B.
'96. On the fracture system of joints. Proc. Boston soc. nat. hist., vol. 27,
pp. 163-183.

Printed, April, 1900.

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h
X

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Boston Society of Natural History.

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APR 20 1900

Prooeedings of the Boaton Society of Natural History.
Vol. 29, No. 11,
pp. 217-222.

A REVISION OF THE SYSTEMATIC NAMES EMPLOYED BY

WRITERS ON THE MORPHOLOGY OF THE

ACMAEIDAE.

B^ M. A. Wii,Lr«»x, Ph.D.

BOSTON:
PRINTED FOR THE SOCIETY.

- Ai'itiL, UK)0.

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k^^'^tm

No. 11. — A Jievision of the Systematic Names employed by
Writers on the Morphology of the Acmaeidae.

By M. a. Willcox, Ph. D.,
Professor of Zoology in Wellesley College, Wellesley, Mass.

In view of the small extent of our knowledge of the Acroaeidae,
it would seem especially desirable that what is known should be
rendered available by the employment of uniform terminology.
This is, however, no less an unattained ideal in this small group
than in other parts of the animal kingdom. It is the purpose of
this brief paper to so arrange and compare the systematic names
employed by various authors as to enable the reader to orientate
himself with the least possible delay.

From 1758 down to the early part of this century all true limpets
appear to have been included under the generic name of Patella,
About 1830 several investigators independently separated off from
the remaining members of this genus a group differentiated by the
possession of a cervical gill, or ctenidium. Eschscholtz, '30,^ called
the new genus Acmaea\ Audouin and Milne Edwards (Cuvier,
'30, p. 326) named it Tect%ire\ Gray, '33, p. 800, termed it Lottia.
This name was obviously of later date than the others, and Gray
himself, '47, p. 158, abandoned it in favor of that introduced by
Audouin and Milne Edwards, which he seems to have Latinized
into Tectura,^ As to the use of Acmaea or Tecturay scientists are
divided. The French hold to the latter, while the rest of the
zoological world has agreed upon Acmaea, I have not found in

1 This reference, which I have been unable to verify, I take from Watson, '86, p. 29.
DaU, '71, p. 237, quotes the same work with the date of Dorpat, 1828. As he gives the
name in its English form, as in a later paper, *78, p. 342, he states that the ** English
reprint," which was published in the spring of 1830, was dated by the author Dorpat,
Jan. 7, 1828, and, Anally, as Watson states that he has made unavaUing search for any
publication to which DalVs reference could refer, I am led to the belief that Eschscholtz's
description did hot issue from the press before 1830. This, however, would not aifect
the question of priority, since, as Watson points out, the Ann. des sci. nat., t. 21, con-
taining as it does reports of meetings held as late as Dec. 13, 1830, could not have issued
from the press before 1831.

> Dall, 71, p. 239, states that he has failed to ftnd any earlier publication of the Latin
form. I have not been more fortunate. Marschairs " Nomenclator " lists Teciura as
published in 1830 by Audouin and Milne Edwards; but this, so far as the form of the
word goes, is an inaccuracy, due undoubtedly to Marschairs habit of Latinizing names.

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218 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

French writers any attempt at a justification of this practice, which
would seem to have arisen from the fact that Tectura, being pro-
posed — at least in its French fonn — at about the same time as
Acf/i/xea, secured a position in the land of its authors which has
never been successfully disputed, although its rival has undoubted
right of priority. The arguments have been fully stated by Dall
and Watson. Bouvier, '87, p. 22, offers one argument unmentioned
in either of these statements — the great similarity between the
names Acmaea and Actnea, The latter was proposed by Ilartmann
in 1821, but, according to Watson (op. cit.), was abandoned by him
the same year. I have l>een unable to consult Hartmann's paper;
but I am informed on excellent authority that his Acmea is derived
from ^aKfiT^; Acmaea, on the other hand, comes from ^atcfialoi
(Rathke, '33, p. 16), and should therefore stand.

Some twenty years laj-er this group of ctenidium-bearing limpets
was itself subdivided. In 1847 Gray, '47, p. 158, apparently with
some hesitation, separated from the others those which have both a
ctenidium and a branchial cordon, giving to this new genus the
name of Scurria, and restricting the name of Ttctura (z=Lott%a) to
those which lack the brancliial cordon. The name of Lottia Gray
was thus abandoned to be revived after nearly twenty years. At
this time Carpenter, '65, pp. 140-141, erected a new genus for a
limpet provided with both ctenidium and branchial cordon, but
having the latter absent in the region of the head. As it appeared
that this very animal had been figured by Sowerby, '20-'25, vol. 1,
pi. 141, as the first mentioned example, and therefore inferentially as
the type of Lottia Gray, for which Gray himself had mentioned no
type species, the name given to the new genus was of course
Lottia. So that Lottia (xray — or more correctly, Lottia Cpr. ex
Qray — represents but a small part of the original I^ottia Gray.
It should be said also that some at least of the more recent authori-
ties have reckoned Lottia as a subgenus of Scnrrla, so that the
systematic rank of tlie group is a matter upon which opinions
vary. And finally it may be remarked that Carpenter, '60,
p. 3, had already listed this species, though without a description,
under the name of Tecturella (jrandis. This name fell, owing to
preoccupation.

In 1834 Broderip described a new limpe^like shell, giving no
account of the animal, and entitled it Scutella, As this name had
already been employed by Lamarck, Gray, '47, p. 168, replaced it

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WILLCOX: SYNONYMY OF THE ACMAEmAE. 219

by ScuteUina, Gray placed the new genus among the Patellidae ;
but H. and A. Adams, '58, vol. 1, p. 460, included it among the
Tecturidae. This appears to have been done on^ the strength of
their own investigation ; at least they give the earliest figure
which I have been able to find of the animal as distinct from the
shell, and mention, although they do not figure, the gill. Scutellina
retained its position in the Acraaeidae (^ Tecturidae) either as a
genus or as a subgenus (of Acmaea) until Dall, '89, p. 410, after a
careful study of the animal, transferred the genus from the Doco-
glossa to the Rhipidoglossa, — a change which has been approved
by Tryon and by Simroth. It is interesting to note that one of
the points which decided D^l to make this transfer — the fact that
the apex of the shell is posterior instead of anterior — was noted
by Broderip in his original description.

Although Scutellina has thus been removed by most authorities
from the Acmaeidae, it is still occasionally included in that group.
It may, therefore, be worth while to call attention to a brief paper
by Pilsbry, '91, p. 88, in which he points out that /Scutellina, having
been employed in 1841 by Agassiz for a genus of echinoderms, is
preoccupied, and must therefore fall as the name of a molluscan
genus. He suggests as a substitute Phenacolepas,
The Acmaeidae then include the following groups : —

Acmaea Eschscholtz — cervical gill, but no branchial cordon.

Scurria Gray — cervical gill and continuous branchial cordon .

Lottia Cpr. ex Gray — cervical gill and interrupted branchial
cordon.
If now we examine works dealing with the moi*phology of the
family, we find these names often incorrectly used. I enumerate
all the papers known to me which treat, otherwise than incidentally,
the anatomy of any one of the Acmaeidae, noting errors of nomen-
clature where I have found them.

1. Rathke, '33, gives a somewhat full description of the anatomy
of Acmaea.

2. Bouvier, *87, p. 22, gives a brief account of the nervous sys-
tem of Tectura (= Acmaea) testtidinalia.

3. Bernard, *90, pp. ^\l~^'lfi^ deals with two species of Tectura
(= Acmaea) which are thus described (p. 217) : " C'est d'abord la
Tectura (Acmaea) pileopsis, qui diff^re des Patelles par la pre-
sence d'une branchie bipectin^e et la Tectura fontainesi, qui
poss^de a la fois une branchie bipectin^e et des lamelles branchiales
circumpall^ales."

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220 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

If we are to infer from this description that T. pileopsis differs
from PateUa^ not only in the presence of a cervical gill, but also in
the absence of a branchial cordon, it may possibly be identified with
Acmaea pileopsis Q. and G., although this species is recorded by
Tryon as belonging to the New Zealand, Indo-Pacific and Austra-
lian region, while Bernard (p. 217) states that his species came
from Chili.

Tectura fontainesi, however, is obviously incorrectly named, and
i s very probably a JScurria. I have not been able to find trace of
any member of the family bearing this specific name.

Bernard has also investigated Lottia peUuciday which he describes
in the following terms (p. 225) : " Les Lottia ou Patina sont de
petits Patellid^s d^pourvua de branchies proprement dites, mais
munis de lamelles pall^ales et de tentacules lat^raux." This is
obviously Patina pdlucida Linn. I have found no justification for
applying the name of Lottia to this genus.

4. Von Erlanger, '92, p. 604, describes the nephridia of an unde-
termined species of Tectura (^ Acmaea) .

5. Haller, '94, describes four Acmaeidae : Scurria, two species
of Lottia Gray, and Scutellina, If Dallas classification of Scutelr
Una be accepted, it is probable, as has been suggested by Thiele,
that we have here to do with a case of incon*ect determination.
The description (pp. 26-27) shows clearly that the animal (sup-
posing it to belong to a described genus of the Docoglossa) is an
Acmaea.

The two species of Lottia Gray- are also Acmaeas ; one lacks the
branchial cordon altogether ; the other has irregular outgrowths
which Haller (p. 27) regards as the anlagen of the cyclobranch
gills. They probably correspond with A, viridtUa Lam. and A.
scutum Orb.

6. Pelseneer, '91, p. 61, describes the eye of Acmaea testu-
dinalis.

7. Thiele, '92, p. 231, describes the structure of the mantle-edge
in CoUisella (Acmaea) digitalis, CoUisella is one of the sub-
genera of Acmaea, and according to both German and American
rules the subgeneric name, when required, should be placed in
parenthesis and interpolated between generic and specific appella-
tions. The name of Thiele's species should therefore read Acmaea
( CoUisella) digitalis.

8. Willcox, '98, treats the general anatomy of Acmaea fragilis.

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WILLCOX: SYNONYMY OF THE ACMAEIDAE. 221

LITERATURE.

Adams, H. and A.

'58. The genera of recent moUusca. 3 vols. London.
Bernard, F^liz.
'90. Recherches sur les organes pall^ux des gast^opodes prosobranches.
Ann. des sci. nat., (7) t. 9, pp. 89-404, 10 pi.
Bouvier, E. L.

'87. Systtoie nerveux morphologie et classification des gast^ropodes pro-
sobranches. Ann. des sci. nat., (7) t. 3, pp. 1-510, 19 pi.
Broderip, W. J.
'34. Description of a new genus of Gasteropoda. Proc. zool. soc., pt. 2,
pp. 47-49.
Carpenter, P. P.

'60. Checklists of the shells of North America. Smithsonian misc. coll.
'65. Diagnoses de mollusques nouveaux proyenant de Califomie et faisant
partie du mus6e de I'institution Smithsonienne. Journal de conchylio-
logic, t. 13, pp. 129-149.
Cuvier, Georges.

'30. Rapport sur trois mtooires de MM. Victor Audouin et Milne-
Edwards, relatifs aux animaux sans vert^bres des c6tes de la France.
Ann. des sci. nat., t. 21, pp. 317-829.
Dall, W. H.
'71. On the limpets ; with special reference to the species of the west coast
of America and to a more natural classification of the group. Amer.
journ.^of conch., vol. 6, pp. 227-282, 4 pi.
'78. Report on the limpets and chitons of the Alaskan and Arctic regions,
with descriptions of genera and species believed to be new. Proc. U. S.
nat. mus., vol. 1, pp. 281-344.
*89. Report of the mollusca of the Blake. II : Gastropoda and Scaphopoda.
Bull. mus. comp. zool., vol. 18.
Erlanger, R. von.

'92. On the paired nephridia of prosobranchs. Quart, journ. micr. sci.,
vol. 33, pp. 687-623, 2 pi.
Eschscholtz, F.

'30. Appendix Kotzebue's Neue Reise. Weimar.
Oray, J. E.

'33. Some observations on the economy ot molluscous animals and on the
structure of their shells. Phil, trans, roy. soc. of London, vol. 123, pp.
771-819.
'47. A list of the genera of recent mollusca, their synonyma, and types.
Proc. zool. soc., pt. 16, pp. 129-219.

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222 PROCEEDDfGB: BOSTON SOCIETY NATURAL HISTOBY.

Haller, B^la.
'94, Studien tt. dooogloase u. rhlpidoglosse Prosobranchier. Pp. 173, 11 pi.
Leipzig.
Pelseneer, P.
'91. Sur I'oeil de quelques gastropodes. Ann. de la soc. beige de micro-
scople, 1. 16, pp. 69-76.
Pilsbry, H. A.

'91. On the use of the generic name Scutellina. NaatilnB, yoI. 5, pp. 88-89.
Rathke, M. H.
'33. Eschscholtz's zoologischer Atlas. Ftinftes Heft, herauBgegeben yon D.
[ate] Martin Heinrich Rathke. Berlin.
Sowerby, James and George B.

'20-'25, The genera of recent and fossil shells. 2 vols. London.
Thlele, J.

'92. BeitrMge zur Kenntnis d. Mollusken. 2, U. d. MoUoskenschale. Zeitr
schr. wiss. zool., Bd. 55, pp. 220-251.
Watson, Robert Bogg.
'86. Report on the Sci4>hopoda and Gasteropoda collected by H. M. S. Chal-
lenger during the years 1873-1876. Challenger reports, vol. 16. Pp. 766,
60 + 3 pi.
Willcox, M. A.
'90. Zur Anatomie yon Acmaea fragilis Chemnitz. Jen. Zeitschr., Bd. 32,
pp. 411-466, 3 pi.

PrirUed AprUy 1900.

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'^'•

Boston Society of Natural History.

RECENT PUBLICATIONS.

' Memoirs. 4to.

The development, .structure, and affinities of tlie genus Etjtiisetuni. By Edward
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The development of Penilia schmackeri Richard. By Mervin T. Sudler.
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Studies iu Diptera Cyclorhajiha. 1. The Pipunculidae of the United States.
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Notes on the reptiles and amphibians of Intervale, N. II. By Glover M.
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Variation and sexual st»lection in man. By Edwin Tenney Brewster.
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Moniloporidae, a new family of Palaeozoic corals. By Anuvdeus W. Grabaii.
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Studies in the gold-bearing slates of Nova Scotia. By J. Edmund Woodnian.
33 pp., 3 plates. 50 cts.

North American wood frogs. By Reginald Heber Howe, Jr. 6 pp. 10 cts.

Some Ilydroids from Puget Sound. By Gary N Calkins. 36 pp., 6 plates.
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The Odonate genus Macrothemis and its allies. By Philip P. Calvert. 32 pp.,
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( )n the veins of the Wolffian bodies in the pig. By Charles Sedgwick Minot.
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Notes on a Carboniferous boulder train in eastern Massachusetts. By Myron
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The genus Antennaria in New England. By Merritt L. Fernald. 13 pp. 15 cts.

The land mammals of peninsular Florida and the coast region of Georgia. By
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A contribution to the petrography of the Boston Basin. By Theodore G.
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Clymene products sp. nov. By Margaret Lewis 5 pp., 2 plates. 15 cts.

The Harvard geographical models. By W. M. Davis. 20 pp., 4 plates. 25 eta.

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JUN 4 1900

Prooeedlngs of the Boston Society of Natiiral History.
Vol. 29, No. 12,
pp. 223-240.

PROCEEDINGS OF THE ANNUAL MEETING, MAY 2, 1000.

BOSTON:
PHIN TED FOR THE SOCIETY.
May, 1000.

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JUN 4 1900

No. 12. — Proceedings of the Annual Meeting, May 2, 1900.
REPORT OF THE CURATOR, ALPHEUS HYATT.

The most notable event of the past year was the retirement of
the Secretary, Mr. Samuel Henshaw, who resigned in order to accept
a position in the Museum of Comparative Zoology at Cambridge.
His resignation was noticed by a formal resolution, but naturally
this did not allude to his long and efficient services during the time
that he was an assistant in our Museum. This gentleman's name
appears first in the Annual Report of 1876-77 when he took charge
of our insects, and it occupies thereafter an annually increasing
importance in the records of the Museum, until he became general
assistant in 1883. A large part of the time from 1876 to 1883 he
had worked for us without pay or with a merely nominal salary,
and during those years had not only accomplished much for the
insects which were under his charge, but had laid the Society under
obligations for important work upon most of the other collections in
the Museum. From 1883 to his election as Secretary in May, 1892,
Mr. Henshaw fulfilled the duties of general assistant with excep-
tional ability. His active connection with the Museum consequently
has lasted for about twenty-three years, and his labor has been felt
in every department and always greatly to the advantage of the
Society. The Curator is, therefore, very glad to be able to say
that, although his work now lies almost wholly in Cambridge, he
still remains connected with our Museum. He has completed the
admirable cycle of his life with us by consenting to remain in charge
of the insects as a voluntary assistant, thus returning to the position
with which he began so many years ago.

The Society has long been in need of some separate room where
the meetings of the Council could be held, and this year, princi-
pally through the efforts of the President, the northwest basement
room was fitted up for this j)urj>ose and also furnished with a large
blackboard and settees, so that it can be used for meetings of sec-
tions of the Society, if any are formed, or by natural history clubs
or small societies that may find it convenient to meet in our build-
ing. Improvements usually have accomj)anying inconveniences,
and this one obliged the Curator to crowd the collections heretofore
stored in two rooms in the cellar into one room and to make other
changes that are not yet entirely completed.

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224 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

The urgent need of means iiith which to do work has been
insisted on in these reports year after year ever since the first year
of the Curator's connection \i4th the Society, and yet in spite of
this, and often repeated appeals in more public ways, the impression
is still prevalent in Boston that we are not in need of money.
Another impression that needs correction is still more unfavorable
to our progress. This is the prevailing opinion that we are not an
educational institution. In spite of all the lecturing to teachers and
to students and to the public that has taken place systematically
and constantly in this building for thirty years past, and in defiance
of moi'e or less frequent newspaper notices of the kind of work done
here, people at large regard us as a Society whose functions, out-
side of a museum that is open to the public for two days in the
week, are strictly private and for the benefit of members alone.
These false impressions lead wealthy people who are continually
giving in other directions to neglect us and even to say, if requested
to give, that a Society ought to take care of itself. A few years
ago I heard one of the most prominent Boston merchants and lib-
eral givers to the cause of education say the same thing about the
Institute of Technology. His complete conversion to the o]i])osite
side within a few years leads me to hope that a brighter future may
be in store for us. Certainly nothing can be more without justifica-
tion than that an institution like ours, devote<l to the encourage-
ment of research through its publications and to the diffusion of
knowledge throligh its museum and lectures, can be self-supporting.
It might be, if a good proportion of the citizens of Boston would l>e
willing to tax themselves to the amount of our annual dues; but
when only a very minute i)roportion of the citizens is willing to do
this, it is practically absurd to expect us to maintain ourselves by
any such means. At present the money for membei-ship is all used
uj), and in fact is not sufticient to meet our annual ai)])ropriation for
publications alone.

In view of this general neglect of our needs it is pleasant to notice
that this year we have received a donation of ten thousand dollars
that the Rev. R. C. Watei-ston left by his will, to be paid after the
decease of Mrs. Waterston. Mr. Waterston was elected a member
in January, 18G0, and his death occurred Feb. 21, 1898. For about
twenty years he was a helpful member of this Society, and he was
selected to give one of the addresses at the celebration of our fif-
tieth anniversary in 18S0. During the last ten years or so of his
life, his increasing disabilities kept him from very active participa-

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HYATT : REPORT OF THE CURATOR. 225

tion in the affairs of the Society, but he was always ready to assist
in every way that he could, and his last generous action shows an
appreciation which is very grateful, since he was thoroughly con-
versant with what had been done and with the future aims of our
work. The Secretary's Report gives the special puq)ose of this
donation, and it is only necessary for the Curator to add that Mr.
Waterston also left his collection under such conditions that we
were able to take out of it whatever was considered suitable for our
uses. It was a miscellaneous collection, but we found in it a con-
siderable number of natural history specimens that could be used in
the laboratory and a few for our Museum.

The remainder of this report is given under special titles designating
the different departments of the Museum in which work has been
done.

Mineralogy and Geology.

Professor Crosby has continued the work on the general collec-
tion of minerals which was interrupted last spring by poor health.
This consists chiefly in weeding out duplicate specimens, which is
in itself a great improvement of the collection, besides making room
for new material. He has gone over the entire collection in this
way, and will use a part of this material in exchanges. A large
amount of work has been done in preparation of Part 3 of the
Geology of the Boston Basin, which, it is expected, will soon be
ready for distiibution.

Teaching in the Museum.

A lady of Boston, as stated in a former report, carried on this
department for a number of yeai*s, and Mr. Grabau's work as
the lecturer and guide to the collections and in other connected
lines of public instruction was fruitful in results that justified the
hope of making this undertaking a permanency. It was a really
unique and successful effort to make collections effective instru-
ments of instruction in j)lace of the lifeless and comparatively
inefficient assemblage of facts they now are in museums, and would
have met with substantial aid if the good work that was done could
have been made known to the proper persons. The Curator there-
fore, feels that apj^eals for the re-establishment of this department
should not be dropi)ed.

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226 PROCEEDINGS: BOSTON SOCIETY NATURAL fflSTORY.

Dynamical Zoology.

If it were not for the great personal interest taken in thi» depart-
ment by the Curator, the difficulties attending its installation would
have long ago led to its abandonment. Considerable work has
been done in this direction by the Curator and Miss Bryant, but as
usual the results appear to be small, owing to the })eculiar difficul-
ties that have to be encountered in selecting specimens that will
show the relations of organisms to their surroundings.

Synoptic Zoology.

The gratuitous work of Mrs. Sheldon in this department, as noted
in previous Reports, still continues, and this and other aid received
from her makes it ponsible to anticii)ate the final completion of this
unique collection and the text of the Guide. The amount of good
work put into this undertaking will then be in such a form that it
can be understood and properly appreciated. The following sum-
mary gives only the bare facts of what has been done, but does not
convey any idea of the amount of work involved in the study of
the literature and the careful judgment and investigation needed for
the selection of the dra^^nngs and specimens mentioned. The prin-
cipal work of the year has been uj)on the Crustacea, Arachnozoa,
and Myriapoda. The ty})e8 of all of these have been described,
eighty for Crustacea, twenty-two for Arachnozoa, and eight for
nuTiapods. The figures selected for illustrations of structure and
develo])ment are as follows: forty-five for Crustacea, thirty-eight
for Arachnozoa, and five for M\niaj)oda. Eighteen pages of the
text on Arachnozoa have been written. A large amount of work
has also been done upon the Insecta, in comparing different systems
of classification, by the same assistant, who has also selected and
described thirty types of this class and has picked out twenty-^ight
figures to illustrate the fossils, the i)i'imitive fonns, and early stages
of development ; and she has also written fifty-one pages of the
text of the Guide for this class. Forty-eight figures of various
grou})s, coelente rates, ecliinoderms, i)elecy]>ods, cephalopods, and
pteropods, have been completed, and the text of the Guide relating
to these has been revised. A beginning has also been made upon
theVertebrata, four ty})es and ten figures of primitive forms having
been selected and described, and some pages of the text written.

Miss Maitin has spent considerable time in making colored draw-
inijs for this collection, under the direction of 3Irs. Sheldon.

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HYATT: REPORT OF THE CURATOR. 227

Botany.

Fortunately the sickness of Miss Carter as mentioned in my last
Annual Report did not prevent her return to duty during the early
part of 1900; and since then this lady has worked in the Museum
and reporte as follows. The small collection received from the
Boston Museum has been catalogued, labeled, and incorporated in
the Museum. The special labeling of the Lowell collection, begun
some years ago, has been completed. Considerable progress has
been made in the systematic arrangement of the collection of dupli-
cates. Fifty specimens of economic fungi have been received from
Seymour and Earle. Twenty-two persons have been permitted to
consult and study in the herbarium.

Paleontology.

Miss Bryant has unpacked and named and catalogued the Cura-
tor's collection of Anticosti fossils, which have been stored in the
cellar for several years. These specimens had been loaned to the
late Prof. James Hall, but were found, upon being opened, not to
have been labeled while in Albany. The same assistant has also
taken care of a small collection received from the estate of Mr.
Waterston, and has spent some time in the identification of the
corals recently purchased from Mr. G. K. Greene.

MOLLUSCA.

Miss Martin has been occupied mainly in the effort to bring
together all of the collections in this department, which, owing to
the absence of proper facilities for storage, have been hitherto neces-
sarily kept in several different places. The Mollusca room has been
furnished with suitable cases for this purpose, and the work of incor-
porating all the different lots of shells into one single systematic
collection has made some progress. Tliis necessarily involves a
large amount of labor that will probably last for several years. The
same assistant and Miss Bryant have worked over, and placed in vials
and proper boxes, and numbered, the single shells, and have cata-
logued all of the Gasteropoda of the Roper collection. The Cyreni-
dae, the most valuable part of the Roper collection, containing a
considerable number of t\T)es of new species and rare shells, were
brought here by Mrs. Roper on her return from the West, and are

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22H PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

now in our collection. A full report on this collection is deferred
until the whole has been i)ro[)erly catalogued and numbered. The
shelves and specimens in Room K, containing? most of the shells,
and those in the gallerj' adjoininij, have l>een dusted and placed in
as good order as practicable, but this labor is really thrown away
as long as the old cases remain in their present condition, with
loosely fitting doors.

Miss Brj-ant has looked over our special collection of New Eng-
land shells and made a list of the species needed to fill gaps in this
series, and has filled a few of these gaps with shells found elsewhere
in the Museum.

The Curator has continued his work upon the Achatinellidae,
especially the ground shells of this family, and has practically
completed the detailed descriptions in manuscript of all the species
of the genera of this division throughout the whole chain of the
Hawaiian Islands. In pursuance of the plan of this work, applica-
tion has been made for the collections of these ground shells stored
in the principal museums of this country. This part of the work
has so far included only the small collections of the Yale University
Museum and the Smithsonian Institution ; but, as it has been going
on for a short time only, and the progress has been rapid, it is
thought that it will not take many months. The Curator will
undoubtedly be able to enrich the collection by exchanges, and he
has already foun«l some exceedingly rare shells and some distinctly
new species in the two collections so far studied.

Rev. II. W. Peck has very generously placed his collection of
Achatinellidae as a loan in the Society's Building, and his Amastras
and other land shells have been named and described. Mr. Oleson
has withdrawn his collection of Achatinellidae, to offer them for
sale elsewhere. There still remain in the neighborhood of 40,000
shells in this building, and nearly a complete set of all the species
of this family ; so there is sufficient material. Special observations
had also been ma«le upon all of the Oleson shells, and tlie only
loss is in the ability to revise manuscript relating to these from time
to time, an omission which will tell more decidedly upon the value
of Mr. Oleson 's collection than upon the memoir in which they are
mentioned.

CRrSTACEA.

Professor Kingsley has coni}>leted the naming of fifty-three lots
of the Amphipoila loaned to him some years since, and these have

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HYATT : REPORT OF THE CURATOR. 229

been returne<l an<l stored in the collection. Through his kind
assistance fifty-three lots of this 'group from the southern coast of
New England have been picked out and forwarded to Prof, H. C.
Bumpus as a loan to aid in his researches. The work of restoring
fade<l labels in the alcoholic collection has been completed by
Miss Martin.

Fishes and Reptiles.

These departrfients have received some desirable additions from
the collection of the Boston Museum, but unluckily none of these
had any locality labels.

Birds axd Mammals.

In consequence of a fire that affected the upper part of the build-
ing of the Boston Museum in 3[ay of last year, the proprietors
turned over to the Society their remaining specimens of birds and
other vertebrates. The birds were found to have been less injured
by dam})ness, smoke, and insects than had been expected ; and
about one third of them were sufiiciently valuable to be retained,
although none of them had special locality labels. A certain num-
ber fell to pieces and were lost, [)erhaps one per cent, altogether.
Miss Bryant was emj)loyed for some time during the summer in
getting the birds together, securing the labels and storing them in
insect-proof cases. After this was done Mr. Batchelder went over
the entire collection in the autumn, identified the si)ecies, and picked
out all of those that were considered suitable for exhibition.
A considerable number of the other vertebrates were looked over
by the Curator, and a number of these were incorporated in the
laboratory collections, and some few were found to be desirable
additions to the Museum collections.

Mr. Batchehler has added by i)urcliase twenty-eight birds to
the NeAv Englantl collection. Five of these are to re))lace badly
faded specimens, but the remainder represent species or important
plumages hitherto wanting in our collection.

Laroratory.

The room in our basement has been used as in previous years by
the da.sses of the Boston Universitv and the Teachers' School of

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230 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

Science. A nunil>er of new (lia^ams have been made by Miss
Martin. The Curator, assisted by Mr. Coles and Miss Martin, has
rearranged the entu^ collection, a task requiring considerable labor,
owing to the confusion into which all parts of this collection had
fallen during the last ten years.

Remarks.

• An unusually large amount of time has been expended this year
upon miscellaneous work not reporte<l upon above, both by the
asHistants and by the Curator.

The Museum has been visited this year on days other than public
days by 494 pupils representing VI schools.

Teaciieus' School of S( iexce.

Mr. Grabau has continued his work in this department without
remuneration. Five short and three long excursions were made
during the spring of 1899 to the seashores in the neighborhood of
Boston, and to some of the freshwater ponds, for the purpose of
studying and collecting living animals. One of the longer excur-
sions was made to Cuttyhunk, one of the Elizabeth Islands, and
occupied four days. The average attendance on the short trips
was fifteen, and there were seventeen persons present on the Cutty-
hunk trip. The longest excursion was made to Bayville on Line-
kin Bay, Maine. This occupied ten days in July. Here a tem-
porary laboratory was opened, and lectures were given. This class
was divided into sections, and j)ractical instruction in the local
geology and botany was added to the marine zoological work, each
section carrnng on one of these subjects. A numl)er of the inhab-
itants of Ba\'\'ille attended this course, and the botanical section
remained for ten dayH longer after the main body of the class had
gone home. The average attendance on this excursion was fifteen.
The third excursion, of ten days' duration, was made in July to
!Monhegan Island, Maine, under the charge of some of the advanced
students of the Teachers' School of Science, who assumed Mr.
Grabau's duties <luring his absence at the West. The average
attendance on this excursion was twenty-one. No systematic work
was attempted in the autumn, but two excursions were made to
localities in the vicinity of Boston. This decline in the autumn is
the natural })relude to the entire cessation of this work, since Mr.
Grabau will probably not remain in this vicinity after this year.

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HYATT: REPORT OF THE CURATOR. 231

Field lessons in geology were given by Prof. George H. Barton
to the pupils of the Boston Normal School. The small number of
these lessons was due to the prevalence of bad weather. It was
stated last year that these lessons would be given up, owing to the
fact that no remuneration was provided for them by the school
authorities ; but in spite of this Professor Barton, when appealed to
again this spring, found the strength of his resolution weaker than
his sympathy for the need of such instruction by ))upils who were
destined to be teachers. Thus in a rich city like Boston the puj)ils
of its normal school are obliged to ask for gratuitous instruction in
order to get the small amount of geological work that they feel to
be necessary. The course has accordingly begun again with a class
twenty-three in number. Many of these pupils, when they
become teachers, enter the regular classes of the Teachers* School
of Science and take the complete four years' course.

It was mentioned in the last annual report that the Tnistee of
the Lowell Institute had concluded to discontinue the out-of-door
work of the Lowell Institute, and that Professor Barton's field
courses would consequently not receive any further sup])ort from
this fund. This misfortune made it necessary to appeal to other
persons for aid, and luckily a patron was found who sufficiently
appreciated Professor Barton's work to give the necessary amount
for continuing his field lessons for one year. The solidity of Pro-
fessor Barton's work, and the high esteem in which it is held by
teachers, and its fine results, are appreciated by all who are ac-
quainted with the facts, and their cessation would be a serious
drawback to the progress of nature study in the schools. Our most
sincere thanks and those of many teachers are due to the generous
donor who has enabled us to continue this work. Professor
Barton reports upon this part of our work as follows.

The regular course of these field lessons in geology, consisting of
ten, was given in the spring of 1S99, April 2*2 to June 24, inclusive.
The total attendance was 149 ; the average attendance being 28.9 ;
largest number present at one lesson 57, smallest number present
at one lesson, which occurred on a rainy day, 8. There were ten
lessons also given in the autumn of 1899, Sept, 16 to Nov. 18,
inclusive. The total attendance in this course was 276, average
attendance 54.8, largest number present at one time 166, smallest
number present at one time 21. A great improvement on the
conduct of previous courses became practicable this year, which ^-ill
greatly add to their efficiency. A system of examinations was

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232 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

begun with the autumn series of lessons, and was- mtroduced as a
voluntary arrangement for all those who wished to get the greatest
profit from the work. These examinations were taken each week
by twenty-four, who were thus obliged to be in regular attendance^
and at the close of the season a final examination of three hours'
length was given on the Saturday following the last lesson.

The spring course for 1900 has neen begun with an attendance of
45, and will l)e reported upon next year. The ordinary field lessons
occupy only a half day on Saturdays, a few take all day like those
to Marblehea<l, Fitchburg, and Clinton, while those to Iloosac
Tunnel take two or three days each.

Lowell Free Courses.

The four years' course begun by IVofessor Barton last winter has
been continued by a series of sixteen lessons of two hours each, or
thirty-two hours of instruction, on hthology and dynamical geology,,
exclusive of the final examination which occupied three hours more.
They began, Dec. 9, 1899, and ended A])ril 14, 1900. The class
numbered IIS, and the average attendance was remarkably large,
being 99.3. This course was as usual a combination of lectures
with laboratory work. Trays of specimens were provided, and the
teachers were require*! to identify the different rocks used and to be
able to recognize these at sight and also to describe their texture
and composition. The mineral trays contained 20 s}>ecimens each^
and about 2,000 specimens were use<l, representing 40 varieties of
the common rock-fonning minerals. The rock trays containetl 12
specimens each, an«l 14 sets were used, representing 168 varieties.
The entire number of specimens use<l was 8,400, making together
with tlie minerals a total of 10,400 specimens provided for class use.
These specimens are now stored in our basement, in cases provided
for their reception. Examinations were carrie<l on throughout the
term as heretofore, an<l a final examination was also held, and care-
ful records are kept by Professor Baiton of the attendance and
standing of every pupil. Professor Barton expresses himself as
highly pleased with the progress of the class and with the attend-
ance ; but he adds that, so far as he has been able to judge, there is
not a trace of any symi)atliv with their work shown by the author-
ities at the head of our i>ublic schools. Fifty- nine meml)ers of this
class were from Boston, and the balance represented thirty-four
neighboring towns.

Dr. R. W. Greenleaf, who had given the lessons on botany for

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HYATT : REPORT OF THE CURATOR. 233

six years beginning in the winter of 1891-92, wa« obliged to resign
on account of the pressure of his professional engagements. The
Curator greatly regretted this, since Dr. Greenleaf had been a most
successful teacher and his courses were very attractive to the best class
of teachers and productive in other results noted from time to time
in previous Reports. Mr. B. II. Van Vleck continued Dr. Green-
leaf's four-year course, completing the third year with sixteen les-
sons of two hours each, beginning Nov. 18, 1899, and ending
March 24, 1900. The number of persons registered was forty, and
the average attendance thirty-two. Thirty took the examination,
and all passed. Two students were for adequate reasons allowed
to be absent and to have an examination later by Mr. Van Vleck.
The subject was the structure and physiology of algae, and numer-
ous preparations and specimens were used, fully illustrating the
more important morphological and physiological facts which it was
desirable to demonstrate clearly on account of their general bearing
in relation to the higher orders of plants. Mr. Van Vleck was
assisted effectively by the work of Miss Cora H. Clarke ; and through
her kindness the class was able to do its work more advantageously
and to receive pressed mounts to the number of 700, representing
25 genera and thirty species. These were gratefully received by
members of the class, who highly appreciated ^liss Clarke's
generosity.

The Curator gave the last series of lessons in a five years' course,
consisting of twenty-two lessons of two hours each, altogether forty-
four hours of instruction, beginning on the 21 st of October, 1899,
and ending on the 15th of April, 1900. The examination has
not yet been held, having been postponed until the second
Saturday in May. The number of lessons exceeds that of any
previous year, but it was necessary in order to cany out the
plan of the whole course and finish it properly. The subjects
were some of the higher orders of Insecta not finished last winter
and the Vertebrata, ending with a special lesson on man. Thei-e
were forty-eight tickets issued, and the average attendance was
thirty-six. This course was fully illustrated as usual with speci-
mens, and the lessons consisted partly of lectures and partly of
obser\'ations made by the students themselves under the direction
of the Curator, the object being instruction in the broad general
facts of structural and functional relations of different animals, with
only enough systematic w ork to enable the pupils to recognize the
natural relations of the types use<l in the class room.

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234 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

REPORT OF THE SECRETARY AND LIBRARIAN,
CHARLES F. BATCHELDER.

Membership.

During the past year seventeen Corporate Members have been
elected. One Honorary Meml)er, Sir William Dawson, and six
Corresponding Members, Mariano Barcena, Elliott Coues, Sir Wil-
liam H. Flower, Hans B. Geinitz, W. T. Hoffman and Alj)honfie
3iilne-Edward8, have died.

Three Corporate Members have died : Elizabeth R. Cormier,
John C. Jackson and Charles T. White, and one Patron, Edward
Wyman. Two Corporate members have resigned.

The membership of the Society, corrected to May 2, 1900, con-
sists of 9 Honorary, 130 Corresponding, and 429 Corporate Mem-
bers, a total of 568. There are 16 Patrons.

The Corporate members elected during the year and the dates of
their election are as follows : —

Cakes Ames, Oct. 18, 1899.
Mary A. Bowers, Oct. 18, 1899.
John G. Graham, Oct. 18, 1899.
Charles Harrington, Oct. 18, 1899.
James J. Minot, Oct. 18, 1899.
Reginald C. Robbins, Oct. 18, 1899.
R. T. Atkinson, Dec. 20, 1899.
Elizabeth E. Bickford, Dec. 20, 1899.
Henry T. Burr, Dec. 20, 1899.
George A. Hathaway, Dec. 20, 1899.
Clement W. Andrews, Feb. 21, 1900.
Laurence Curtis, Feb. 21, 1900.
George W. W. Dove, Feb. 21, 1900.
W. E. C. Rich, Feb. 21, 1900.
M. Eva WaiTen, Feb, 21, 1900.
Arthur W. Fab-banks, April 18, 1900.
. Flora G. Roper, April IS, 1900.

Meetings.
Fourteen regular meetings of the Society have been held during

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BATCHELDER : REPORT OF SECRETARY AND LIBRARIAN. 235

the year. The attendance has varied from twenty-eight to ninety-
three, the average being about fifty-one.

Nineteen formal communications have been made at the meetings
by fifteen persons, and eight papers have been presented by title.
The lantern has been used to illustrate papers at eleven meetings. >

The meetings, attendance and communications have been as
follows : —

May 3, 1899. Annual meeting. Thirty-four persons present.

Reports of the Curator, Secretary, Librarian, Treasurer, and

Trustees.
Mr. W. L. Tower. A quantitative study of the migration and

variation of the Colorado potato-beetle.
May 17, 1899. General meeting. Thirty-one persons present.
Prof. Alpheus Hyatt. Exhibition of Gage's series of brook

and lake lampreys from New York.
Mr. A. W. Grabau. Evolution of the Fusidae.
November 1, 1899. General meeting. Seventy-seven persons

present.
Prof. W. M. Davis. Geographical notes of a year in Europe.
Dr. H. S. Pratt. The embryonic history of imaginal discs in

the brachyceran Diptera. (By title.)
Mr. Glover M. Allen. The species of Evotomys of eastern

North America. (By title.)
Mr. Arthur M. Edwards. Diatoms of the U. S. geological

survey of the Territories. (By title.)
November 15, 1899. General meeting. Sixty-one persons present.
Mr. J. G. Jack. Forest aspects and problems in central

Colorado.
December^, 1899. General meeting. Ninety-three persons present.
Mr. William Brewster. Nesting habits of some New England

birds.
3Ir. H. T. Burr. The discoveiy of fossils in the Roxbury con-
glomerate.
Dr. R. P. Bigelow. Anatomy and development of Cassiopea

xamachana, (By title.)
December 20, 1899. General meeting. Forty-five persons present.
Mr. A. W. Grabau. Notes on a geological excursion in the

Rocky Mountains of Colorado.
Mr. R. H. Howe, Jr. Description of a new race of Horizopua

virena (Linn.) . (By title.)

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236 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

Jaiiuary 3, 1900. (Teneral meeting. Fifty-six persons present.

Prof. F. G. Wright. New light on the age of the Niagara

gorge.
Mr. R. T. Young. A brief report on the mammals of Prince
Edward Island. (By title.)
January 17, 1900. (xeneral meeting. Thirty j>erson8 present.

Dr. R. T. Jackson. Some cases of oUl age characters in inver-
tebrates.
February 7, 1900. General meeting. 'Thirty persons present.

Dr. C. S. Minot. Notes and illustrations of mammalian devel-
opment.
Dr. G. H. Parker. The correlation between the size of litters

and the number of mammar}' glands in the swine.
Dr. R. P. Bigelow. Notes on the development of Cassiopea,
Dr. C. S. Minot. On the solid or closed condition of the

intestine in the chick.
Dr. C. S. Minot. On a hitherto unrecognized form of blood
circulation without capillaries, in the organs of vertebrata.
(By title.)
Prof. M. A. Willcox. A revision of the systematic names
employed by writers on the morphology of the Acmaeidae.
(By title.)
February 21, 1900. (xeneral meeting. Forty-five })ersons present.

Prof. C. II. Fernald. The g\l>sy moth in America.
March 7, 1900. General meeting. Fifty-eight }>erson8 present.

Dr. Frank Russell. The Moki snake dance.
March 21, 1900. General meeting. Eighty-six persons present.
Prof. W. M. Davis. Glacial erosion in the Alps and in
Norway.
April 4, 1900. General meeting. Twenty-eight j)ersons present.
Dr. G. n. Parker. The neurone theory in the light of recent
investigations.
April 18, 1900. General meeting. Thirty-seven }>ersons present.
Mr. J. II. Enierton. The common species of American spiders.

PUBLK ATIONS.

During the year the following publications have been issued : —
Proceedings of the annual meeting, May 8, 1899. Proceedings,
vol. 29, no. 1, 43 pp.

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BATCHELDER : REPORT OF SECRETARY AND LIBRARIAN. 237

Variation and Bexual selection in man. By Edwin Tenney
Brewster. Proceedings, vol. 29, no. 2, 17 pp.

Notes on the rejitiles and amphibians of Intervale, New Hamp-
shire. By Glover M. Allen. Proceedings, vol. 29, no. 3, 13 pp.

Studies in Diptera Cyclorhapha. 1. The Pipunculidae of the
United States. By Garry de N. Hough. Proceedings, vol. 29, no.
4,10 pp.

Contributions from the Gray Herbarium of Harvard University.
New Series. — No. 17. By B. L. Robinson and J. M. Greenman.
Proceedings, vol. 29, no. 5, 22 pp.

The development of Penilia achmackeri Richard. By Memn
T. Sudler. Proceedings, vol. 29, no. 6, 23 pp., 3 pis.

List of marine mollusca of Coldspring Harbor, Long Island, with
descriptions of one new genus and two new species of nudibranchs.
By Francis Noyes Balch. Proceedings, vol. 29, no. 7, 30 pp., 1 pi.

The blood vessels of the heart in Carcharias, Raja, and Amia.
By G. H. Parker and Frederica K. Davis. Proceedings, vol. 29,
no. 8, 16 pp., 3 pis.

The occurrence of fossils in the Roxburj' conglomerate. By
Henry T. Burr and Robert E. Burke. Proceedings, vol. 29, no.
9, 6 pp., 1 pi., 2 cuts.

On a hitherto unrecognized form of blood circulation without
capillaries in the organs of vertebrata. By Charles Sedgwick
Minot, LL.D. Proceedings, vol. 29, no. 10, 31 pp., 12 cuts.

A revision of the systematic names emi)loyed by writers on the
morphologA^ of the Acmaeidae. By M. A. Willcox, Ph.D. Pro-
ceedings, vol. 29, no. 11, 6 i)p.

In connection with the publications mention should be made of
the bequest of 810,000, received this year from the late Robert C.
Waterston, the income of which is to be devoted to the Society's
publications.

Library.

The additions to the library have been : —

8vo. 4to. Folio. Total.

Volumes
Parts
Pamphlets
Maps

Total

3S8

408

lS-21

355

2177

4G7

499

tioTG

461

'31H4

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238 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

The library contains 25,629 volumes, 1401 current, or otherwise
incomplete, volumes, and 13,311 pamphlets.

Among the more important gifts to the library during the year
have been the bequest of the late Robert C, Waterston (120 vol-
umes) , some books from Mr. James M. Barnard (29 volumes) , and
Field's Bibliography, received from the Trustees of the Thompson
Fund.

New exchanges have been arranged with the Philosophical Society
of Washington, the Kongliga Universitets-Biblioteket, Upsala, Swe-
den, the Royal Geographical Society of Australasia, and the Bemice
Pauahi Bishop Museum, Honolulu.

One exchange has ceased. The Society now exchanges its pub-
lications with 435 institutions and periodicals.

Eight hundred and ninety-two books have been borrowed by 182
persons ; 441 have been borrowed for use in the building ; the
library has been consulted about 350 times.

Four hundred and one volumes have been bound in 270 covers.

Twelve volumes of the Proceedings of the U. S. National Mu-
seum have been indexed. Current volumes of serials pre\nously
indexed, are indexed as received.

Walker Prizes.

The subjects for competition appointed for 1900 were: —

1. Stratigraphy and correlation of the sedimentary formations of
any part of New England.

2. A study in palaeozoic stratigraphy and correlation.
The Committee has reported the following awards :

A prize of one hundred dollars for the essay entitled, " A study
in palaeozoic stratigraphy and correlation : the Hudson River beds
of the neighborhood of Albany and their taxonomic equivalents,"
by Rudolf Ruedemann, Ph. D.

A i)rize of fifty dollars for the essay entitled '* Cephalopod zones
in the Carboniferous of Xoith America : a study in interregional
coiTclation,'* by James Perrin Smith.

The subject for the award in May, 1901, is: —

A monograph on any problem connected with, .or any group
belonging to, the Xoith American fauna or flora.

Digitized by VjOOQ IC

BOUVE : REPORT OF TREASURER.

239

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240 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

The reportH of the Trustees and of the Auditing Committee were
presented, and it was voted to a(x*.ept the several reports.

The Society then proceeded to ballot for officers for 1900-1901.
Messrs. J. H. Blake and J. B. Woodworth were appointed to collect
and count the ballots. They reported the election of

PRESIDENT,

CHARLES SEDGWICK MINOT.

VICE-PRESIDENTS,

CHARLES p. BOWDITCH, HENEY W. HAYNES.

WILLIAM G. FARLOW.

CURATOR,

ALPHEUS HYATT.

SECRETARY,

CHARLES F. BATCHELDER.

TREASURER,

EDWARD T. BOUVfe.

LIBRARIAN,

CHARLES F. BATCHELDER.

COUNCILLOR FOR ONE YEAR,

Mis8 Clara E. Cummings.

COUNCILLORS FOR THREE TEARS,

S. L. AimoT, Mi.sH Catharine I. Ireland,

William S. Bryant, Benjamin Joy Jeffries,

William M. Davis, N. T. Kidder,

Samuel Henshaw, William H. Niles.

Printed, May, l-^oo.

Digitized by VjOOQ IC

Boston Society of Natural History.

RECENT PUBLICATIONS.

Mkmuiuh. 4t(>.

The development, structure, and affiaities of the p'nus E(}ui8etum. By Edward

C. Jeffrey. 30 pp., 5 plates. $1.(X).
Localized staples in development in plants and animals. By Robert T. Jack.son.

65 pp., 10 plates. $2.00.
raocKEDiNOs. 8vo.
A revision of tlie systematic names employed by writers on the morphology' of

the Acmaeidae. By M. A. Willcox. 0 pp. 10 cts.

On a hitherto unreco^niiz*Hl fonn of blood cifculation without capillaries in the
organs oi veruebrata. By Charles Sedgwick Minot. 31 pp. 36 cts.

The occurrence of fos^^ils in the Roxbury con.^lomerate. By Henry T. Burr

and Robert K. Burke. 0 pp., 1 plate. 20 cts.
Tae blood vessels of the heart in Carcharias, Raja, and Amia. By G. H. Parker

and F. K. Davis. 10 pp., 3 plates. 26 cts.
List of marine mollusca of Coldspring Harbor, Long Island, with d'CHcriptions

of one new genus and two new sjxjcie.s of nudibranchs. By Francis

Noyes Balch. 30 pp., 1 plate. 36 cts.
Tiie development of Penilia schmackeri Richard. By Mervin T. Sudler.

23 pp., 3 plates. 30 cts.
(\»ntributi{)ns from tlie (»ray herbarium of Harvard university. New series,

no. 17. 1. Revision ot the genus (iymnoiomia. 2. Supplementary' not^s

uimn Calea, Tridax, and Mikania. By B. L. Robuison and J. M.

Ciieenman. 22 pp. 26 cts.
Studies in Diptera Cyclorhapha. 1. 'Hie Pipunculidae of the United States.

By Garry de N. Hr)ugh. 10 pp. 10 cts.
Notes on the reptiles and amphibians of Intervale, N: H. By Glover M.

Alien. 13 pp. 16 cts.
Viiriation and st^xual selection in man. By Edwin Tenney Brewster.

17 pp. 26 cts.
MoniloporidcU% a new family of Palaeozoic corals. By Amadeus W. Grabau.

l(i pp., 4 plates. 26 cts.
Studies in the gold-bearing slates of Nova Scotia. By J. Edmund Woodman.

33 pp., 3 plates. 60 cts.
North American wood -frogs. By lieginald Heber Howe, Jr. 6 pp. 10 cts.

Some Hydroids from Puget Sound. By Gary N Calkins. 36 pp., 0 plates.

60 CIS.
The Odonate genus Macrothemis and its allies. By Philip P. Calvert. 32 pp.,

2 plates. 60 cts.
On the veins of the Wolffian bodies in the pig. By (Miarles Sedgwick Minot.

10 pp., 1 plate. 26 cts.
Notes on a (Carboniferous boulder train in eastern Massachusetts. By Myron

L. Fuller. 14 pp. 16 cts.
The genus Antennaria in New England. By Merritt L. Fernald. 13 pp. 16 cts.
The laud mammals of peninsular Florida and the coast region of Georgia. By

Outram Bangs. 70 pp. 76 cts.
A contribution to the petrography of the Boston Basin. By Theodore G.

White. 40 pp., 6 plates. 06 cts.
Clymene producta sp. nov. By Margaret Lewis. 6 pp., 2 plates. 16 cts.
The Harvard geographical models. By W. M. Davis. 20 pp., 4 plates. 25 cts.
The role of water in growth. By C. B. Davenport. 12 pp. 16 cts.

Digiti

ized by Google

Prooeadln^ Of the Boston Sooietj of Natural History.

Vol. 29, No. 13,
pp. 241-272.

JUL 17 ISM

THE EMBRYONIC HISTORY OF IMAGINAL DISCS IN MELOPHAGUS

OVINUS L., TOGETHER WITII AN ACCOUNT OF THE

EARLIER STA(;ES IN THE DEVELOPMENT

OF THE INSECT.

By H. S. Pratt., Ph. D.

WlTH^SEVKN I'LATKS.

BOSTON:
PRINTED FOR THE SOCIETY.

JlTNE, 1900.

Digitized by VjOOQ IC

JUL 17 1900

No. 13. — The Embryonic History of Imaginal Discs in Melo-
phagHfS ovinus X., together with aii Account of the Earlier
Stages in the Development of the Insect}

By H. S. Pratt, Ph.D.
With seven plates,

CONTENTS.

Page

Introduction 241

Historical :

1. Imaginal Discs in the Larva and Pupa . ... 242

2. Imaginal Discs in the Embryo 248

The Earlier Developmental Stages of Melophagus :

1. The Development of the Egg to the Completion of the Blasto-

derm 251

2. The Formation of the Mesoderm and of the Proctodeum and

Stomodeum 255

The Origin of the Imaginal Discs :

1. The Cephalic Discs 259

a. The Early Development of these Discs . . . • 259

6. The Involution of the Head 264

2. The Thoracic Discs 267

3. The Discs of the External Genitalia 268

4. The Discs of the Internal Organs 269

Methods 269

Bibliography 270

Introduction.

In 1897 I published in " Psyche " a preliminary account of the
origin and early development in Melophagus ovinus of those funda-
ments of the imaginal head, wings, and appendages, which may be
present in the larva of holometabolic insects and are known as
imaginal discs. The present paper is an extended account of the
same matter.

Melophagus belongs to the small group of brachycerous Diptera

1 Contributions from the Zoological Laboratory of the Museum of Comparative Zoology
£X Harvard College, under the direction of £. L. Mark, No. 111.

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242 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

called the Pupipara, which are closely related to the muscids.
Leuckart (*S7) early showed the striking similarity of structure of
the larval and pupal forms of the Pupipara to those of the muscids ;
in a former paper ('93) I have emphasized the same fact ; MOggen-
burg ('92) has homologized the mouth parts of all the pupipars with
the fly's proboscis ; Brauer ('85) has shown that the Pupipara are
degenerate flies and has placed them next to the Muscidae in his
classification of the Diptera.

The Muscidae are classical objects for the study of imaginal
discs. It was with them that Weismann inaugurated the study
of these interesting bodies, and, since the appearance of his first
paper in 1803, Ganin, Kowalevsky, Van Rees, and many others
have publidied the results of investigations of them. All of these
studies, however, have been on the development of the discs during
the larval and pupal period of the insect's life. The embryonic
history of the discs, which includes their origin and the first stages
of growth, has not been studied ; and no positive information exists
concerning them except a short statement contained in a paper by
Graber ('89), which will be spoken of later on, and the preliminary
paper by myself, mentioned above.

Historical.

1. Imagimd Discs tfi the Larva and Pupa. — Swammerdam
(1737-38) was the earliest investigator to observe that fundaments of
the imaginal thoracic appendages in the higher insects, and even of
the head in some cases, do not appear first in the pupa, but are present
in the larva. lie was thus the first observer of imaginal discs. He
showed that in the larva of Culex, Apis, and Pieris the fundamentd^
of all the legs and wings lie beneath the thoracic integument. From
his time down to the present generation no additions were
made to the knowledge of the subject. Numerous investigators,
however, made observations similar to those of Swammerdam.
Lyonnet (1760) described and figured the two pairs of imaginal discs
in the dorsal portion of the meso- and metathoracic segments of the
caterpillar, and added the supposition that they were the funda-
ments of wings. Herold ('15) described the same discs, and cor-
rectly interpreted them. Burmeister ('35) also very accurately
described the imaginal wing-discs of the caterpillar, as did Louis

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PRATT: IMAGINAL DISCS. 243

Agassiz ('51) somewhat later. Leuckart ('57) mentions the tho-
racic and cephalic imaginal discs of Melophagus, but without describ-
ing them or knowing their ultimate fate.

It was in the years 1863 to 1866 that Weismann ('63, '64, '66)
laid the foundations of our present knowledge of imaginal discs in
a series of investigations on the development of Musca and other
Diptera, both brachycerous and nematocerous. He ('63, p. 229) found
that the dipterous larva, which is apodous, and in the case of the
Brachycera, acephalic, contains within its thorax six pairs of disc-like
bodies. In Simulia, a nematocerous dipter, which is the first form he
studied, they are situated near the animal's integument, although
apparently not in connection with it. There are three pairs ven-
trally located, each pair belonging to one of the thoracic segments,
and three pairs dorsally located, these being similarly distributed
among the thoracic segments. All of these bodies he found in close
relation to nerves or tracheae or both. He further found that they
remain function less during the life of the larva, although increas-
ing greatly in size with the growth of the larva, and that during the
metamorphosis they develop into certain organs of the imago.
He called them, consequently, imaginal discs. He found that the
three pairs of ventral discs develop into the imaginal legs, the
dorsal metathoracic pair into the balancers, the dorsal mesothoracic
pair into the wings, and the dorsal prothoracic pair into the anterior
pupal spiracles, when these are present.

In Corethra (Weismann, '66), also a nematocerous dipter, he
found similar conditions.

In Musca he ('63, '64) found the conditions very much more
complicated. The six pairs of discs just mentioned he found pres-
ent ; but, instead of being located near the integument, they were
sunk into the centre of the animal's body. An additional pair of
discs was also i)resent in the forward portion of the thorax, directly
in front of and closely applied to the brain -garfglia ; these he found
were destined to develop into the imaginal head. Weismann also
found that only a small portion of the larval body passes directly
into the imaginal body, the greater part of it undergoing disintegra-
tion, so that the tissues entirely lose their identity, and afterwards
the imaginal body is built up anew from the imaginal discs. To
this process, the entire significance of which, however, was not
understood until later, he gave the name " histolysis P

These early papers of Weismann have furnished the starting-

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244 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

point for a large number of investigations upon the imaginal
discs. He confined his studies to the Diptera ; other insect
groups were soon investigated. Landois (*71) following Herold,
Agassiz, and most of the older authors, studied the imaginal wing-
discs to be found beneath the dorsal' thoracic integument of the
caterpillar. Ktinckel d'llerculais (*75) showed that in the larva of
Volucella, a muscid, the imaginal discs, although situated at a dis-
tance from the integument, are connected with it by a delicate
chord, the remnant of an invagination. He also discovered two
pairs of imaginal discs near the hinder end of the body of the larva,
which develop into the external genital organs.

Ganin (*76) studied imaginal discs in several groups of insects, —
namely, the Hymenoptera, Neuroptera, Coleoptera, Lepidoptera,
and Diptera, — in the larvae of all of which he found wing-discs, and
in those which are apodous, leg-discs as well. He added to these
observations the discovery, in the larvae of brachycerous Diptera, of
other discs than those described by Weismann. That author
believed that the hypodermis of the larval abdomen went directly
with qiodifications to form that of the imago. Ganin now showed
that in, and forming a part of, the hypodermis of each of the eight
abdominal segments of the muscidian larva, are four discs, two
dorsal and two ventral, the tissue of which resembles that of the
thoracic discs, and that they form the starting point for the grovrth
of the imaginal hypodermis of the abdomen. Ganin likewise discov-
ered similar discs in the epithelium of the larval mid-gut, whose fate
it is to form in the same way the imaginal mid-gut ; and he also dis-
covered the important fact that each imaginal disc is made up of
two kinds of embryonic tissue, ectoderm and mesoderm. Further-
more he discovered the amoeboid mesoderm cells which destroy the
larval organs during histolysis.

Dewitz ('78) took up the study of imaginal discs for the purpose
of reviewing the work of Ganin, Landois, and Weismann. A few
years later Viallanes ('82) studied afresh the post-embryonic devel-
opment of the Muscidae, and laid the basis of our present knowledge
of the histological details of the process of histolysis in the meta-
morphosis of insects.

In 1882 and 1883 Metschnikoff published the first of his epoch-
making studies on the destruction of tissues in certain invertebrates
by leucocytes, or, as he called them, phagocytes. He discussed
Ganin's observations and especially that of the destruction of the

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PRATT: IMAGINAL DISCS. 245

larval organs by amoeboid mesoderm cells during histolysis in the
pupal muscid. These cells, he suggests, are none other than phago-
cytes. Shortly afterward. Van Rees ('84) and Kowalevsky ('85)
proved the correctness of this position. They showed that the
process of histolysis consists of the ingestion and digestion of the
functional larval tissues by phagocytes, and the building up of
imaginal tissues from imaginal discs.

Kowalevsty ('87) in another paper took up the investigation of
the histolysis of the internal organs of Musca where Ganin and
Viallanes had left it, and gave the first complete account of these
processes. He showed the exact method by which the muscles,
digestive tract, and hypodermis of the lai*va are destroyed by
phagocytes and the imaginal organs reconstructed from imaginal
discs. And in the following year Van Rees ('88) published his
extensive paper on the post-embryonic development of muscids, and
completed our knowledge of this phenomenon. He showed that
when the muscidian larva enters upon the pupal stage, histolysis is
inaugurated by the destruction of the larval muscles, which become
unfunctional du-ectly after pupation and a natural prey to the
phagocytes. Soon the thoracic hypodermis and the inner organs
are a^ttacked, and at the same time the imaginal discs begin to grow
and widen out, supplying the place of the tissues which are being
destroyed. The continuity of the hypodermis and of most of the
internal organs is thus at no time broken, an obsei-vation which
Kowalevsky ('87, p. 585) also made, correcting at the same time the
statement to the contrary made by Viallanes ('82, p. 221) . As these
processes go on, the two large cephalic imaginal discs, which form
two irregularly shaped sacs extending as diverticula from the
dorsal wall of the pharynx back to the brain, begin to move for-
ward, dragging the brain with them. Their anterior ends bend and
pass ventrally, embracing the pharynx between them. At the same
time their communications with the pharynx enlarge and their
lumina fuse more and more completely with the pharyngeal lumen
until they meet in the median line and form one single median
opening, which, ever increasing in size, finally extends the entire
length of the discs. The lumina of the discs and of the pharynx
thus become completely merged and form together a single con-
tinuous space, and the walls of the discs and of the pharynx a single
continuous vesicle. This is the head vesicle or " Kopfblase," which
is destined to become the imaginal head. This vesicle remains

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246 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

buried within the pupal thorax Until near the end of the pupal
period, when it evaginates and forms the completed head of the
insect. This evagination has been obsei'ved by Weismann ('64,
p. 259) to be the consequence of the pressure of blood, which at the
right moment is forced from the abdomen into the thorax and pushes
the head vesicle forward.

The metamorphosis of the thorax goes on simultaneously with
the formation of the head vesicle. In proportion as the larval
hypoderrais disappears under the attacks of the phagocytes, as
already mentioned, the edges of the imaginal discs grow and
take its place, forming the imaginal hypodermis. As we have
seen, there are six pairs of these discs, a doi-sal and a ventral
pair being present in each thoracic segment. They lie at a dis-
tance from the integument, near the centre of the larva. Each
disc is, however, connected by a very fine hollow cord with that
portion of the hypodermis of the segment to which it genetically
belongs, and where it is destined to appear as an extremity. This
cord, first discovered by Ktlnckel d'llerculais, gradually shortens,
and its lumen enlarges. The disc is thus brought nearer the
surface, and as it advances it increases in size. The lumen of the
cord then opens through the hypodermis, and the cord itself fin-
ally becomes so wide and short that the disc is brought through
the hypodermis to the outside. The hollow cord is of course
obliterated by this process, and the edges of the proximal end
of the disc are brought into direct contact with the hypodermis.
The disc has now assumed its position as an extremity. It is an
appendage of the body-wall ; it has become irregularly cylindrical
in shape, and possessed of a number of constrictions and folds,
which in the case of the ventral discs are equivalent to the joints of
the future leg.

The metamoq^hosis of the abdomen is retarded ; it does not
begin until that of the head and thorax is well advanced. Then
in each abdominal segment the two ventral and four dorsal discs
(Van Rees found two additional dorsal discs in each segment)
begin to grow and take the place of the disappearing larval
hypodermis.

In a paper of my own ('93), which contains the results of a study
of the larva of Melophagus ovinus, is contained a full description
of the imaginal discs of this insect. Melophagus, being a pupipar,
and closely allied to the muscids, we should expect to find the same

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PRATT: IMAGINAL DISCS. 247

imaginal discs in its larva as in the muscidian larva. And we do, in-

dac. prHhz.
dsc. ms'thx.

dsc. mVthx.

A B

A, ventrcU frontal section ; B, dorsal frontal section ; dsc. pr^thx., dsc. msHhx.^
and dsc. mVthx.^ pro-, meso-, and metathoracic imaginal discs.

deed, find similar conditions in general ; but there are some interest-
ing and instructive differences. The larva is apodous and acepha-
lous, like the muscidian, but in many ways it is less highly special-

d 0 » 0 0 0 olo e

oooooooooo

ized ; indeed, it seems, in some respects, to occupy an intermediate
position between Corethra and Musca. In the position of the
thoracic discs, for instance, it closely resem-
bles Corethra. These discs are found just
beneath the integument in two very regular
rows, and not near the centre of the larva,
as in the muscidian larva. The accompany-
ing cuts represent frontal sections through
the anterior end of an old larva (Fig. A being
a ventral and Fig. B a dorsal section), show-
ing the position of the thoracic discs. In struc-
ture the meso- and metathoracic discs stand
exactly halfway between the same discs in Co-
rethra and in Musca. In Corethra, according
to Weismann ('66, p. 78), all the thoracic
discs are of larval origin, and each is a
double fold of the hypodermis, of which it
remains a part, as is shown by Figure C.
In Melophagus, on the other hand, these
discs arise in the embryo; they are also

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248 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

double folds of the hypodermis, but become constricted off from
It, as shown in Figure D. In Musca the discs are also of embryonic
origin. They also become constricted off from the hypodermis;
but, instead of remaining where they have originated, they suffer
removal toward the centre of the larva, as is shown in Figure'E,
and the peripodal membrane, as Van Rees calls the outer wall of
the disc, lengthens to form the hollow cord which connects it with
it3 old position at the hypodermis.

In the cephalic discs the conditions are similar to those in Musca,
but even more complicated. Instead of a single pair of head-discs
there are two pairs present, one dorsal and one ventral. The dorsal
pair corresponds to the muscidian head-discs in every respect ; they
are destined to form the dorsal and lateral portion of the imi^inal
head together with the compound eyes. The ventral head-discs
have no counterpart in Musca. In the embryo they arise as a
single median thickening, from which paired diverticula develop.
From the bottom of each of these diverticula, which in the larva
project from the ventral pharyngeal wall, there extends into its
lumen a long projection. The diverticula fuse in the median line
during the latter portion of the larval period, and the wall thus
formed between them gradually disappears, so that in the full-grown
larva the ventral discs appear as a single ventral diverticulum of the
pharynx, at the bottom of which a pair of long projections extends
towards the wide opening. The fate of these discs is to form the
ventral portion of the head, the paired projections being the funda-
ments of the proboscis. The formation of the head- vesicle during
the metamorphosis proceeds in a way similar to that in Musca.
The ventral disc fuses early at its lateral edges with the dorsal pair;
the communications between both ventral and dorsal discs and the
pharynx become rapidly larger (in the old larva they have already
become very wide), and soon the discs and the pharynx form
together a single vesicle, the head-vesicle.

2. Imaginal Discs in the Embryo. — As already stated, the em-
bryonic history of imaginal discs has not been studied in the higher
insects. The following are the speculations of authors as to their
origin.

In Corethra, a nematocerous dipteron, Weisraann ('66) found that
the imaginal discs do not make their appearance until after the last
larval moult. In the Brachycera, on the other hand, he found the
cephalic and thoracic discs present in the youngest larvae, and he

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PRATT: IMAGINAL DISCS. 249

concluded that they must arise in the embryo. Concerning, however,
the exact manner in which they take their origin, he was not able
to make any positive statements, but was of the opinion that they
first appear within the body-cavity as proliferations of the epithelial
coverings of tracheae and of the neurilemma of nerves. In Core-
thra, on the other hand, he observed that the discs arise as folds
of the larval hypodermis, and he calls attention to the fundamen-
tally different method of origin of imaginal discs in these represent-
atives of the two great divisions of the Diptera.

Landob ('71) , investigating the development of wings in the larva
of Lepidoptera, follows Weismann in his conclusions as to their ori-
gin in the embryo. Ganin ('76) also supports Weismann as to the
embryonic origin of imaginal discs in all the different groups of
insects he studied. KUnckel d'Herculais ('75) was the first one who
discredited Weismann 's conclusions. He found each disc in the
muscidian larva to be connected with the hypodermis by means of
a cord, and he rightly concluded that it resulted from an invagina-
tion and that the discs are therefore of ectodermic origin. Weis-
mann (*64, p. 139) also, it is true, saw these connecting cords, but
failed to interpret them correctly. Dewitz ('78, '81) after a study
of lepidopterous larva came to the same conclusion concerning the
origin of the imaginal discs as Kttnckel d*Herculais, namely, that
they are ectodermal invaginations. Pancritius ('84), also studying
the Lepidoptera, reached the same conclusion. Balfour, in his text-
book ('80) declared that, notwithstanding the authority of Weis-
mann to the contrary, the cephalic and thoracic imaginal discs of
Musca must be derivatives of the ectoderm, as they are in Corethra.
Kowalevskjr ('86), after a study of the embryo of Musca, declared
himself unable to determine the method of origin of the imaginal
discs. He arrived at the negative result, however, that they do not
arise as growths from the epithelium of tracheae, but that tracheae
and nerves unite with them while they are still young.

Van Rees ('88) in his studies of the muscidian larva, demonstrated
the existence of a fine lumen, a continuation of the peripodal space,
in the cord connecting the disc with the hypodermis, and showed
that both lumen and peripodal space are lined with a fine cuti-
cula. He asserts that this discovery is anatomical proof that the
cord, the peripodal membrane, and the disc itself have all been
parts of a single invagination of the embryonic ectoderm in exactly
the same way as the imaginal discs in Corethra arise as in vagina-

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250 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

tions of the larval hypodermis. Graber ('77- '79), however, thought
otherwise. In his valuable treatise ** Die Insekten," part 2, p. 563,
following the results of the first paper of Dewitz ('78;, which had
just been published, he introduces a diagram (Fig. 206, E) repre-
senting the thoracic discs taking their rise as deep invaginations of
the hypodermis. lie does not, however, commit himself to this
interpretation, but says " Eine solche Annahme entbehrt aber
yorlaufig jeglicher BegrUndung, und so dtirfte es doch besser sein,
das Vorkommen einer endogenen Insek ten-Metamorphose einfach
anzuerkennen als sie mit Gewalt zu einem Vorgang umzudeuteln
der mit den bisherigen Beobachtungen nicht Ubereinstimmt."
Shortly after the publication of the above-mentioned paper of Van
Rees, Graber ('89) published an extensive series of studies on the
embryonic development of the Muscidae and certain other insects,
in which the origin of the imaginal discs is touched upon. He
adheres to the conclusions of his former work just quoted, and
takes exception to those of Van Rees. Supported by certain obser-
vations on the embryonic development of Lucilia, he asserts that
the anatomical evidence of that author does not prove the cephalic
and thoracic discs to be hypodermal invaginations. Graber's obser-
vations were made on the cephalic discs and were as follows : The
first traces of these discs which he found were a pair of thick plates
composed each of a siiiffle layer of epithelium situated at the right
and left of the pharynx of the larva and in connection with its lat-
eral walls (see his Figs. 116, 117, 117*). The origin of these
plates, therefore, he did not observe, but the fact that when he first
noticed them they were i7iter7ial structures, and the further fact
that they were not at this early stage in the form of sacs, but of
plates, led him to think that the sac-form these discs possess in their
later and larval developmental stages is a secondary adaptation.
In other words, he believed these discs to arise in the body-cavity
of the animal as epithelial thickenings which afterward assume a
sac-form. He asserts also that the possibility cannot be denied that
the thoracic imaginal discs have a similar origin. In fact, this is
the interpretation of the matter he himself believes.

In later years, Verson ('90) and Gonin ('94) , after studying Lepi-
doptera, Bugnion ('91), after studying a hymenopteron, and Wahl
('99), a dipteron (Eristalis) , have adopted the opinion of Van
Rees, that imaginal discs are of ectodermal origin^

My own investigations on the embryology of Melophagus will

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PRATT: IMAGINAL DISCS. 251

show that Graber's conclusions were in great part false. I shall
show that in the Brachycera the imaginal discs of the head and
thorax all arise in the embryo first as thickenings of the ectodermic
body-wall, and not in the form of endogenic plates; that these
thickenings soon invaginate and form simple pockets in the ecto-
derm. I shall show that the cephalic invaginations are formed
before the involution of the embryonic head characteristic of
brachycerous development, and that as a result of this involution,
whereby the so-called pharynx of the larva is formed from the walls
of the embryonic head, the discs come to form diverticula of the
pharynx, in which condition they are found during the entire larval
and the first part of the pupal period. It will also be shown that
the thoracic invaginations finally become separated from the embry-
onic ectoderm, and form the thoracic discs as we find them in the
larva; also that the external genital organs take their origin in
the form of imaginal discs in the same way as do the thoracic
extremities.

The Earlier Developmental Stages of Melophagus.

1. The Development of the Egg to the Completion of the Blasto-
derm,, — The formation of the blastoderm in the embryos of brachy-
cerous Diptera has been observed and described by the following
authors, — in Musca vomitoria by Weismann (*64, '82) , Kowal-
ovsky ('86), Blochmann ('87), Ilenking ('88), Voeltzkow ('89), and
Graber ('89) , and in LuciUa by Graber (*89). The accounts differ
considerably from one another, although they are descriptions of the
same process and all the authors mentioned are well-known, expe-
rienced, and competent observers. On this account, the details of
blastoderm formation in Melophagus will be a useful contribution
to the subject.

Melophagus is not, however, a favorable object for the study
of the earlier stages of insect development, because of the impossi-
bility of obtaining considerable numbers of its eggs and of deter-
mining the age of the embryos within them. The female insect
produces only one egg at a time and at intervals of several
weeks; each egg must be dissected from the maternal uterus,
where development goes on, and the age of the embryo can be
ascertained only after the egg has been properly stained, and then

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252 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

only approximately by comparing it with other embryos. I should
not, therefore, have selected the eggs of Melophagas for the inves-
tigation of this phase of insect development. My problem, as the
title indicates, was quite a different one, the study of imaginal discs ;
but while searching for embryos containing these structures, I
collected a considerable number illustrating all the different phases
of the animal's development.

The egg of Melophagus has almost exactly the shape of that of
Musca, as observed by Weismann ('64) and Blochmann ('87). It b
an elongated cylindrical object, tapering at the poles, being blunter
at the posterior than the anterior end, with ^ length of 1.2 mm.
and an average breadth of 0.3 mm. at its widest part. As seen from
the side, its ventral outline is convex, its dorsal outline slightly con-
cave. It is covered by a two-layered chorion, the outer layer being
much thicker than the inner one, and by a delicate vitelline mem-
brane. As it lies in the maternal uterus the main body-axes of thfe
developing embryo correspond to those of the mother's body. The
micropyle forms a large funnel-shaped depression in the anterior
end of the chorion (PI. 1, Fig. 4), which sinks deeply into the
yolk, and the deep dent it makes deforms the anterior end of the
developing embryo until near the termination of the embryonic
period. The micropyle is always found filled with a dense mass of
spermatozoa.

The very young uterine egg has also a structure similar to that of
the egg of Musca, but differs from it in one important particular. It
consists of a web of granular protoplasm within which lies a mass of
spherical yolk granules, but the peripheral layer of clear protoplasm
is exceedingly thin (PI. 1, Fig. 1). It has thus no " Keimhaut-
blastem," as Weismann has called the thick peripheral layer of clear
protoplasm first found by him in the muscine egg. This is rather
remarkable, as the presence of the Keirahautblastem is characteris-
tic of the higher insects, having been demonstrated in Musca by
Weismann ('64) and others, in the Coleoptera by Heider (*89) and
Wheeler ('89), in the Lepidoptera by Bobretzky ('78;, in the
Hymenoptera by Grassi ('84) , and in the Hemiptera by Witlaczil
('84), and the absence of it is also characteristic of the lower insects^
as shown by Ayers ('84), Heymons ('95), Wheeler ('89), and Korot-
neff ('85) in the Orthoptera, and by Brandt ('69) in the Pseu-
doneuroptera. The only other instance with which I am acquainted
of the failure of this layer in one of the higher insects is in Lucilia^

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PRATT: IMAGINAL DISCS. 253

a muscid, in which, according to Graber ('89), the hinder portion of
the egg is provided with it, while the forward portion lacks it.
Graber does not find even the thinnest peripheral protoplasmic
layer in the forward half of the egg of this fly, but asserts that the
yolk-balls abut immediately on the vitelline membrane, — an obser-
vation the accuracy of which seems to me on general principles
extremely doubtful.

Melophagus, however, as will be seen, acquires a Keimhautblas-
tem before the completion of the blastoderm.

The processes leading to the maturation and fertilization of the
egg were not observed. The segmentation-nuclei were first ob-
served forming an irregular group near the centre of the egg.
Figure 1 (PI. 1) represents a section of an egg in which about ten
of these nuclei were counted, all of which were migrating through
the yolk towards the periphery. Around each nucleus is a
zone of clear protoplasm, possibly the result of the absorption of
yolk granules near it. Each nucleus, moreover, as it moves toward
the surface, leaves behind it a path of clear protoplasm. Only four
of these nuclei are seen in the section, but portions of the paths of
others are visible. It is necessary to call attention to the irregular
nature of the migration of these nuclei. In Musca the earlier seg-
mentation-nuclei are described by Blochmann and Voeltzkow ('89)
as advancing in very regular order towards the surface of the egg
from the centre and arranging themselves in positions approxi-
mately parallel to the surface as they advance. Kowalevsky (*86)
observed that the segmentation-nuclei in Musca arrive at the sur-
face of the egg first in the hinder portion, then in the forward,
and lastly in the central portion. Voeltzkow ('89) observed in the
same animal that at all portions of the egg they arrive at the same
time. In Melophagus, as will be seen in Figure 1, most of the
nuclei are near the centre of the egg, but are advancing quite irreg-
ularly towards the surface, as is indicated by their plasma-paths.
The penpheral plasma-layer in this egg is very thin, being but 5 ^
thick.

Figure 2 shows a section of an egg in which blastoderm formation
has advanced much farther. The paths of the advancing nuclei
form here a network among the yolk granules. Some of the nuclei
are seen to be in the process of division ; but the great majority of
them are spherical bodies with distinct nuclear membranes and few
chromatin granules. Many have reached the periphery of the eggy

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254 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

and it will be readily seen that there is not the slightest order in
their arrangement, nor is there any regularity in the way they
advance. Figure 3 represents a portion of the same blastoderm on
a larger scale. The peripheral protoplasmic layer has the same
thickness as in the last egg mentioned, except at those places where
nuclei have entered it. Here its thickness is increased by the
diameter of the nucleus, which on the average is 12.5 /i.

Figure 4 represents a section of an egg in which the formation
of the blastoderm is still farther advanced. The nuclei begin to
crowd one another at the surface of the egg, and, not finding room
for themselves in the narrow, peripheral, protoplasmic layer, they
begin to project outward, and thus form protuberances on the
outer surface of the egg, giving a section the wavy outline shown
in the figure. This protoplasmic layer has increased somewhat in
thickness (probably as a result of the consumption of yolk granules
by the rapidly dividing nuclei), and is now 7.5^ thick between the
nuclei. As will be seen in Figure 5, which shows a much enlarged
view of a portion of this blastoderm, the nuclei have nearly the
same diameter as those shown in Figures 2 and 3, and are prolifer-
ating much more rapidly. The protoplasmic paths are not so
numerous as in the egg last described, and will become less
frequent from now on, showing that the yolk granules move
together again, thus obliterating the paths. The gradual thickening
of the peripheral protoplasmic layer also probably goes on at the
expense of these paths.

These processes all continue. The segmentation nuclei at the
surface increase very rapidly in number, and begin to decrease in
size as they become more and more numerous, and the protuber-
ances which they occupy begin to crowd one another (PI. 1, Fig. 6).
These do not, however, show a tendency to merge with one another.
Each protuberance becomes rather more clearly defined and dis-
tinct, and its sides, from forming acute angles with the plane of the
former surface of the egg, as shown in Figure 4, finally come to
form right angles with it (Fig. 6) . The surface of the egg then
presents a curious appearance, being like the pavement of a street
the blocks of which are separated by deep narrow spaces. The
average diameter of the nuclei in the egg represented in Figure 6
is 5 /A. The peripheral protoplasmic layer has also increased in
thickness, and now measures 15 ^ at points between the nuclei.
It now constitutes a blastema, such as is present in Musca before
segmentation begins.

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PRATT: IMAGINAL DISCS. 255

The nuclear paths in this egg have entirely disappeared. Their
disappearance undoubtedly accounts largely for the increase in the
thickness of the blastema.

The steps in the formation of the blastoderm between this stage
and the completed blastoderm I have not observed, but I believe
the concluding process to be as follows : The nuclei and their pro-
tuberances still further increase in number, and the former decrease
in size ; the latter are brought into immediate contact with one an-
other and their walls fuse and become the lateral boundaries of the
future blastodei-m cells; these boundaries are carried still farther
toward the inner surface of the blastema, and the cells are finally
completed by the ^ formation of a wall bounding their inner
ends. In the completed blastoderm (PL 1, Fig. 7) there is still a
narrow blastema present, and the diameter of the nuclei has fallen
to 2.5 IX. No inner or secondary blastema, such as is described in
Musca by Graber ('89) and other authors, is present. I did not
observe the formation of the pole-cells.

2. The JFormation of the Mesoderm and of the Proctodetim and
Stomodeam, — At the time of its completion, the blastoderm is
composed of narrow cells of equal height throughout. In the
centre of the ^^o^ are numbers of so-called yolk-nuclei. Inasmuch
as these yolk-nuclei are homologous to the primitive endoderm in the
gastrulation of the majority of animals, as has been determined by
Heymons (*95, *97) and other authors, the stage of development in
the ontogeny of Melophagus in which the blastoderm is completed,
as represented by Figure 7, would be the gastrula-stage. The next
step in the development is the formation of the germinal plate and
the median mesodermal band. The cells on the concave (dorsal)
side of the ^^'g diminish somewhat in height (PI. 2, Fig. 8); those
on the convex (ventral) side rapidly proliferate along the median
line ; a slight depression appears in the blastodeim along the mid-
ventral line, on the inner surface of which a ridge of cells, the
primitive mesoderm {cr8,ms\lrm.)^ is raised, projecting into the
yolk. The ectodermic cells of this ventral region become elongated,
and with the mesoderraic cells constitute the germinal plate. The
plate does not, however, confine itself to this portion of the egg, but,
as is common in the Diptera, quickly extends itself to portions of the
dorsal side, the thickened ectodermic portion encircling both poles
of the egg and occupying about a third of the dorsal surface at each
end (PI. 2, Fig. 9). The primitive mesodermal plate also extends

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256 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

beneath these dorsal ectodermic thickenings, but is not formed at
the same time at the two poles of the egg. At the anterior end it
extends dorsally more rapidly than at the posterior portions (Fig. 9) .
I found no trace of the lateral gastrular invaginations which Graber
('89) describes in Calliphora as accompanying the median invagi-
nation, and which, as Korschelt and Ileider ('92, p. 812) have
remarked, undoubtedly mark the edges of the germinal plate.

The germinal plate, as in other Diptera, is superficial in position
and unprotected by embryonic envelopes, the amnion being repre-
sented by only the merest rudiments at the anterior and posterior
extremities of the germinal plate (PI. 2, Fig. 9, am.), on the dorsal
surface of the egg. These rudiments mark the limits of the head-
fold and the tail-fold, the former being the dorsal portion of the
germinal plate at the anterior end of the egg {pli, ce,) , and the latter
the dorsal portion of the germinal plate at the posterior end of the
egg {pli, ca.) . Even these rudiments of an amnion disappear in the
stages immediately following.

Figure 9 represents a sagittal section of an embryo somewhat
older than that shown in Figu^:e 8. The anterior end of the e^'g
is marked by the presence of the micropyle (riir^jyy,) . The head-
fold (pli, ce.) extends as far as the anterior amniotic ru^liment and
contains mesodenn, whereas the tail-fold (pli, ca.), which extends
to the posterior amniotic rudiment, consists as yet exclusively of a
thickened ectodermic plate.

The proctodeum and stomodeum both make their appearance as
ectodermal depressions on the dorsal poition of the germinal plate.
In common with Musca, as observed by Graber ('89) and Voeltzkow
(*89) , the proctodeum appears first. Figure 10 (PI. 2) represents
a sagittal section of an embryo somewhat older than the one previ-
ously mentioned; in it the proctodeum has the form of a deep
invagination in the tail-fold, while the stomodeum has not yet
appeared. The point in the head-fold where the stomodeal invagi-
nation will appear is, however, plainly shown in the thickening near
the end of the fold (eras, stmd,). It will also be noticed in this
embryo that the primitive mesoderm has extended beneath the
tail-fold, that the posterior amniotic rudiment has disappeared, and
that the germinal plate has begun to show traces of segmentation.
Tracheal invaginations have not yet made their appearance.

Figure 12 (PI. 2) shows a stage still older than the one just
discussed, in which both proctodeum and stomodeum are present.

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PRATT; IMAGINAL DISCS. 257

The proctodeal invagination {prd.) has become much deeper, and
at its inner end the beginnings of the Malpighian tubules have
made their appearance (tb, mpg,). The stomodeal invagination
(stnuL) forms a deep depression in the head-fold a third of a milli-
meter from the anterior end of the egg, A surface view of the
dorsal aspect of an egg in this stage of development is represented
in Figure 11 (PI. 2), and one of the lateral aspect, in Figure 13.
In both these figures the head-fold (pit. ce,) is seen to be sharply
marked off from the remainder of the dorsal integument by the
presence of the rudiment of the amnion (am). The tail-fold is not
marked off with the same distinctness for two reasons : the posterior
rudiment of the amnion, which appeared in Figure 9, has now dis-
appeared ; moreover the fold itself does not occupy as much of the
dorsal surface of the egg as it did in the previous stages, as is shown
by the position of the proctodeum nearer the posterior pole of the

The primitive mesoderm also shows a marked advance in develop-
ment in eggs of this age. The band or ridge of mesoderm which
took its origin in the mid-ventral line of the germinal plate has sep-
arated into two lateral bands which, as in other insects, occupy
positions to the right and left of the mid-ventral line. Figure 13
shows these bands (tae. 7n8\lrm) as they appear in a lateral view of
the embryo ; Figure 15 {tae. ms'drm) /in a cross-section. It will be
seen that the mesodermal cells have entirely abandoned the mid-
ventral portion of the egg, a 8[)ace being left between the yolk and
the ectoderm which represents the fundament of the coelom (coel.).
The formation of these lateral mesodermal bands, as just described,
takes place, however, only in the middle portions of the egg. At
the two ends of the egg and in the head- and tail-folds the meso-
derm does not entirely abandon the median portions of the egg,
although it expands into the lateral portions of it. The mesoderm
of the ventral side is continuous at the ends of the egg with that of
the head- and tail-folds. Thus at each end of the egg there is a
continuous layer of mesoderm lining the entire inner surface of the
ectoderm (Fig. 12), instead of two lateral bands, such as are found
in the middle portions of the egg.

In embryos of this stage segmentation is distinctly indicated in
the ventral portions of the germinal plate, ten or eleven segments
being represented (PI. 2, Figs. 12, 13). The tracheal invagina-
tions also appear (Fig. 13), there being eleven pairs of them, of

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258 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

which the first two pairs are near the anterior pole of the egg. The
endoderm has now made its appearance and is in process of surround-
ing the yolk. As has been demonstrated by Voeltzkow (*89) and
Graber ('89) in Musca, by Heymons ('95) in Forficula and various
orthopterous insects, and by L^^caillon ('96) in Coleoptera, so in
Melophagus the endoderm is a derivative of the inner ends of the
stomodeal and proctodeal invaginations. The first indication of it
is a proliferation of cells at the inner end of the proctodeum. This
extends itself forward, in the form of a single layer of epithelium,
along the dorsal surface of the yolk (the proctodeum having a
dorsal position) to the head-fold (Fig. 12), and also around the
sides of the yolk toward the ventral side of the egg (Fig. 15,
en^lrm,). As is to be seen in Figure 10 (prd,)^ the proctodeal
invagination, when it first appears, is bounded on its inner end by
the mesodermic layer of the tail-fold. The boundary between
mesoderm and ectoderm in this region is always perfectly easy to
determine, because of the very different character of the cells of the
two germ-layers. As the proctodeal invagination increases in
depth and the fundaments of the Malpighian tubules begin to
appear, the mesodermic layer gradually becomes thmner, until it
entirely disappears. The cells of the anterior (deep) portion of the
ectodermic invagination then proliferate rapidly and give rise to
endoderm (Fig. 12, en\lrm.).

The proctodeum, since it appears earlier than the stomodeum,
apparently gives rise to the greater part of the endoderm, — an
observation which was also made by Graber ('89) on Musea. In
the stage represented by Figure 12 the endodermal epithelium
has extended forward from the proctodenn to the head -fold, and
lies beneath the stomodeum, but without having fused with it. I
do not believe, however, that the stomodeum fails to cooperate in
the production of the endoderm. As shown in the figure last men-
tioned, the stomodeum has just pushed its way through the meso-
dermic layer, which at first bounds its inner surface ; in the stages
succeeding this, as will be shown very soon, this inner surface fuses
with the endoderm and apparently aids in its formation.

No trace of coelomic sacs appears in the mesodertn at any
time.

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PRATT: IMAGINAL DISCS. 259

The Origin of the Imaginal Discs.

1. The Cephalic Discs: — a. The Early Developnient of these
Discs. — Soon after the stage of development represented by Figures
11-13 (PI. 2), but before the lateral mesodermal bands have
extended to the dorsal side of the embryo and joined each other in
the mid-dorsal line, and before the endoderm has extended around
from the dorsal to the ventral side of the embryo (Fig. 15, e7i^drr)i),
those imaginal discs which are destined to form the imaginal head
have made their appearance. Three crescentic thickenings of the
ectoderm appear in the vicinity of and partly encircling the stomo-
deum, a median one in front of the stomodeum, and two paired ones
behind it (PL 3, Fig. 17). The median thickening (dsc, ce. m,) is
destined to form the ventral cephalic disc, which during the meta-
morphosis develops into the ventral portions of the imaginal head
together with the mouth-parts ; the paired thickenings (dsc, ce,)
form the dorsal cephalic discs, which are homologous with the
cephalic discs of the Muscidae, as described by Weismann, Van
Rees, and others. These develop during the metamorphosis into
the dorsal and lateral portions of the imaginal head. The median
disc has no homologue in the Muscidae.

There is no period in the development of the embryo of Melopha-
gus when a distinct head is present, although it seems probable that
the head-fold, since it forms the forward end of the germinal band,
represents the head. It never shows, however, the slightest trace
of segmentation. In Musca, according to Weismann ('64) the head
is, at a certain stage of the embryo's development, quite as distinctly
segmented as the trunk. As will be seen in Weismann's Figure 71,
the embryo is divided into fourteen or fifteen segments, of which
eleven belong to the body, and the remainder to the head. The
anterior portion of the germinal band does not as in Melophagus,
extend onto the dorsal side of the egg, the stomodeum heing sub-
terminal. As the development of the embryonic fly advances, the
cephalic portion gradually becomes reduced in size relatively, and
its segmentation becomes less distinct. When the young larva is
finally born its head has been reduced to a mere rudiment at the
forward end of the body, without a trace of segmentation, and is
smaller than any of the body segments. Consequently, of the
twelve segments composing the fly larva the first alone represents
the head.

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260 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

In the development of the embryonic Melophagus the head-fold
has a history similar to that of these cephalic segments of Mosca.
As the embryo advances in its development the head-fold diminishes
in size, and the stomodeum, which first appeared at some distance
from the anterior end of the egg on the head-fold, migrates forward
towards its definitive position at the anterior end, probably as the
result of a longitudinal concentration or shortening of the germ-
band. The history of the tail-fold is the same. It also becomes
reduced in size as the proctodeum migrates towards the posterior
end of the embryo. This migration b much more rapid than that of
the head-fold. Thus, when the young Melophagus larva is finally
born, it resembles externally in a marked degree the new- bom fly
larva. It consists of twelve segments, of which eleven belong to the
body, and one, which represents the rudiment of the head-fold, to
the head ; the head- and tail-folds have disappeared, the germ-band
having entirely retreated from the dorsal side of the embryo.

The further development of the cephalic discs goes on band in
hand with this migration of the stomodeum towards the anterior
pole of the egg. The paired discs develop much more rapidly than
the median one. The latter remains a mere thickening of the ecto-
derm, without showing signs of invagination, during the entire
migration of the stomodeum, so that when this organ has reached
its final position at the anterior end of the egg the median disc
has simply changed its position. It is now an ectodermal thicken-
ing on the ventral side of the embryo just beneath the mouth
opening (PI. 3, Fig. 22, and PI. 4, Fig. 29, dac. ce. m.).

The paired discs, on the contrary, early in the course of their
movement forward begin to invaginate. The convex margin of
each of the crescentic thickenings becomes much thicker than the
concave, and along this outer (convex) margin a groove-like invagi-
nation is formed (PI. 2, Fig. 14, dsc. ce.). At the stage of devel-
opment represented by Figure 14, the right and left halves of
the germinal band in the region of the body have not proceeded
more than half way towards the dorsal side of the eggy as will be
seen in Figure 15 (tae, ms'dmi)^ which represents a cross-section
through the middle of the same embryo as the one shown in Fig-
ure 14. The endoderm has not yet extended to the ventral side of
the yolk (Figs. 14, 15, en'drm,)^ the tracheal invaginations form
deep sacs (Figs. 14, 16, i'vag, tr,), and the nervous system has not
yet made its appearance.

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PRATT: IMAGINAL DISCS. 261

An embryo which has advanced somewhat beyond the stage
just discussed is illustrated in Figures 18-21 (PI. 3), which repre-
sent sections through a single animal. The stomodeum (Fig. 18,
8tmd,) has become,a deep invagination, which is situated on the dor-
sal side of the embryo, 0.16 mm. from its anterior end. It has thus
moved 0.17 mm. nearer the anterior end than it was at the stage
represented in Figure 12. The inner end of the invagination has
now fused with the endoderm. The forward ends of the paired
cephalic discs appear in this section (Fig. 18, dec, ce,). At a distance
of 0.08 mm. back of this section is the section represented by Figure
19, which shows these discs where the invaginations are deepest.
The invaginations are here long, curved, deep slits, one on each
side of the stomodeum, which have a diagonal position in the head-
fold, the median end of each being directed posteriorly. It will
also be obsei-ved that in this embryo the endoderm has completely
enclosed the yolk. At the anterior end of the embryo it has fused
with the stomodeum (Fig. 18), while at the posterior end it is seen
to be continuous with the proctodeum (Fig. 21, en'drm.). The nerv-
ous system has made its appearance in the form of a pair of longi-
tudinal cords of cells on the ventral wall of the body, one on each
side of a mid-ventral ectodermal ridge (PI. 3, Figs. 18-20, n, v.).
In the head-fold a longitudinal nerve-cord makes its appearance
immediately beneath each cephalic invagination and on the median
side of the slit-like invagination (Figs. 18, 19, g7i, cb.) ; these con-
nect with the ventral nerve-cord at the forward end of the embryo.
These nervous fundaments in the head-fold are undoubtedly the
beginnings of the cerebral ganglia. This early establishment of a
relation between the brain of the animal and the paired cephalic
discs is important, since the relation becomes more intimate as
development advances. In the larva and pupa the cephalic discs
are so firmly joined to the cerebral ganglia that Weismann ('64),
who first discovered the corresponding discs in Musca, gave them
the name " Hirnanhange."

A pair of large spherical bodies, apparently of nervous tissue,
appears in the head-fold at this stage of development (PI. 3, Fig. 18,
ffn, ala.) at the right and left of the stomodeum, and near the for-
ward ends of the cephalic invaginations. The origin of these very
noticeable bodies I did not observe ; but they arose between the
stages represented by Figures 14 and 18, and probably from the
ectoderm. They are at this stage quite unconnected with any other

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262 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

organs. It is very difficult to identify them satbfactorily, but ap-
parently they belong to the pharyngeal (sympathetic) ganglionic
system, and I have called them ganylia allata (see Heymons, 'dS,
p. 49), inasmuch as they do not arise from the dorsal stomodeal wall,
but probably from the outer ectoderm, whence they migrate dorsally
to the position they finally occupy. In other insects in which these
or at least similar ganglia appear, they originate as thickenings or
invaginations of the ventral body-wall in the neighborhood of the
maxillae, and migrate dorsally either to fuse with each other in the
mid-dorsal line just above the stomodeum (Forficula) or to remain
separate (Gryllus). In Melophagus, since the head-fold and the
stomodeum occupy the dorsal portion of the germinal band, these
ganglia probably do not have a ventral, but rather a lateral, origm
(PI. 3, Fig. 18, gn, ala.) , But, as will be seen in subsequent stages,
they migrate to a position immediately beneath the dorsal body-
wall, fuse together, and enter into relations with important pharyn-
geal organs.

No neuroblasts appear in any part of the nervous system at
this early stage of development, the cells being spherical bodies
of nearly equal size. The germinal band has not, in the stage just
discussed, grown over the dorsal side of the animal (except, of
course, on the head- and tail- folds) to complete the formation of the
back.

In the next stage the stomodeum (PL 3, Fig. 22, atmd,) has
reached the forward pole of the body. The cephalic discs have
also changed their positions and migrated farther foi-ward. The
median disc is now situated immediately ventrad of the mouth
opening {dsc. ce. 7n.) ; the paired discs have moved still farther
forward, and are now 0.15 mm. from the anterior end of the
animal ; but their essential character has not changed. Figure 23
(PI. 4) represents a parasagittal section of the same embryo of
which Figure 22 represents a slightly oblique sagittal section. It,
together with Figure 26, — which represents a cross-section of an
embryo of the same age, — shows the relation of the paired discs to
the surrounding organs. These discs (dsc. ce.) are seen to be no
longer dorsal to the embryonic intestine, as in the previous stages,
but to have moved to a position in front of it.

The nervous system has developed considerably. The paired
nerve-cords (n. v.) have fused with the mid- ventral ectodermic
ridge, and neuroblasts appear throughout their entire extent. The

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PRATT: IMAGINAL DISCS. 263

paired cerebral ganglionic fundaments {gn, cb,) beneath the paired
discs have increased in size. Figure 23 shows their connection with
the ventral cords. The ganglia allata ([/n, ala.) have moved to the
dorsal side of the embryo, where their anterior ends have fused
with each other, and occupy a median position (PI. 3, Fig. 22, and
PI. 4, Fig. 26, gn, ala.)^ while their paired posterior portions lie
back of the paired discs (Fig. 28, gn. ala.), A still better idea of
the position of these bodies is given in Figures 43 and 44 (PI. 7),
although these represent sections of a much older embryo, Figure
44 (gn. ala.) showing the posterior portions of the ganglia, and
Figure 43 (g7^. ala.) their juncture. Their anterior fused portion,
even in the earlier stage (PI. 3, Fig. 22, gn, ala.) , has much elon-
gated, and extends forward to a mass of mesodermic tissue, dorsal
to the forward portion of the stomodeum, which is fast developing
into a group of muscle-libres (Ing.), whose later history will be
found to be interesting. In the dorsal wall of the stomodeum is to
be seen a slight evagination (gn./., PL 3, Fig. 22; PI. 4, Fig. 26),
which I take to be the fundament of the ganglion frontale. It is a
very transitory structure, soon disappearing without leaving a trace
in the later history of the animal.

Segmentation has become much more strongly marked in the
forward portion of the embryo, but has almost disappeared from the
hinder portion of it. In Figure 22, which represents a somewhat
oblique sagittal section, we see three deep grooves in the ventral
ectoderm, which mark the boundaries between head, prothorax,
mesothorax, and metathorax, respectively. Two following shallow
grooves mark the boundaries of the metathorax, the first, and the
second abdominal segments respectively.

The arrival of the stomodeum at the anterior end has transformed
that part of the body of the embryo. The mouth is a transverse,
slit-like opening (PI. 4, Figs. 25 and 21jStnid.). Its ventral lip
is the median cephalic disc, which, however, is still only a thick-
ened portion of the ectoderm. Its dorsal lip is a conical struc-
ture, and projects freely forward above it (PI. 3, Fig. 22 ; PI. 4,
Fig. 27, big.). This structure is of great importance in the lar-
val life of the insect, for it acts as a sucking tongue by means of
which the animal ingests its milk-like food. (For a description
of it in the larva, see Pratt, '93.) When it first appears, this organ,
as will be seen in Figure 22, forms the dorsal wall of the anterior
end of the stomodeum, and its anterior tip projects beyond the

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264 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

remainder of the body. These relations are also to be seen in
figure 27 and in the cross-sections shown in Figures 24 and 25, Fig-
ure 24 being a section through the extreme anterior end of the ani-
mal, in which the ectoderm is the only tissue cut, and Figure 25 being
a section just back of it. The sucking tongue is composed of two
distinct tissues, an outer epithelium and /an inner muscular core
(Figs. 22, 25, Ing.), At the stage represented by these figures the
muscle-fibres are just beginning to differentiate. The hinder sur-
face of this muscle mass is in direct contact with the fused median
portions of the ganglia allata (Fig. 22). Figures 27 and 28 repre-
sent surface views of an embryo in this stage of development.

It will be noticed in Figure 22 that a communication between
the intestine and the proctodeum is established. The growth of the
proctodeum has thus advanced faster than that of the stomodeum,
as no communication has yet developed between the stomodeum and
the forward end of the intestine. The opening between the mid-
intestine and the end-intestine is, however, closed again before the
birth of the larva and remains so during the larval life of the animal,
a peculiarity of structure which the larva of Melophagus shares
with that of the honey-bee.

In the animal represented by these figures, the germinal band
has extended over the dorsal side, and the back is thus closed.
The heart is present in the form of a delicate tube extending from
the ganglia allata to the hinder end of the body.

In embryos in this stage of development (and not in any other)
a pair of deep invaginations appears in the outer ectoderm below
the mouth (PI. 4, Figs. 24, 25 and 27, gl, saL); these are probably
fundaments of the salivary glands. They are entirely transitory
structures.

b. The Involution of tlie Head. — An important change now
takes place in the develoj)ment of the for\Yard end of the body.
The paired discs, which hitherto have consisted of two ectodermic
thickenings embracing each a simple, diagonally placed slit (PI. 4,
Fig. 26, else, ce.), now move forward and towards the median line,
where the slits finally meet and unite, forming thus a single, trans-
verse, slit-like opening which extends nearly across the embryo. It*
is only the upper or ectal portions of the invaginations, however,
which thus fuse ; their inner or ental portions increase very much
in volume and depth, though remaining free from each other,
and extend posteriorly in the body-cavity to the vicinity of the

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PRATT: IMAGINAL DISCS. 265

cerebral ganglia, with which they lie in contact. The paired
discs have thus become a single structure with a single, median,
dorsal, anterior opening, backward from which extends an irregu-
larly Y-shaped invagination. Figure 30 (PI. 5), which represents
a sagittal section, shows the median opening and the median por-
tion of the invagination. Figure 31, which represents a parasag-
ittal section, shows one of the branches of the Y-shaped invagina-
tion (dec. ce.) ; and Figure 32, a parasagittal section laterad of the
one last mentioned, shows the irregular structure of the disc.

The ventral median disc (Fig. 30, dsc. ce. m.) has not changed
its position or character, except to become much thicker.

A comparison of the structures of the embryo represented in
Figure 22 (PI. 3) with those of the one represented in Figure 29
(PI. 4) , which is the same as that of the last three figures dis-
cussed, shows that the development in the latter has been consid-
erable. The muscle fibres of the sucking tongue are distinctly
developed, but the nerve which proceeds from its base to the ganglia
allata, and, also, these ganglia themselves, have not changed their
character. The paired ventral nerve-cords and the cerebral gan-
glionic fundaments (Fig. 31) have developed considerably, the
former having fused with the median ectodermic ridge, which has
now effected a separation from the ectoderm, and thus come to
form with the paired nerves a single structure. Neuroblasts are
present throughout the entire extent of both ventral and cerebral
nerve-masses, but are not present in the ganglia allata nor in the
median nerve proceeding from it. It will also be noticed that the
anus has shifted its position from the hinder end of the animal '
to the ventral side near the hinder end of the raid-intestine, the
position it occupies in the larva. A communication has also
appeared between the storaodeum and the intestine (Fig. 30) .

The involution of the head of the embryo now takes place. An
ectodermic fold starts back of the cephalic discs, both dorsally and
ventrally (PI. 5, Fig. 30), and grows rapidly forward towards and
over the mouth. The mouth, together with the ventral disc {dsc,
ce. m.) just below it, the muscular tongue {Ing.)^ and the common
opening of the dorsal discs (of. m.) just above it, is rolled in by
this process. A new mouth is thus formed (PI. 6, Fig. 34, or,),
and back of it a new portion of the digestive tract (phy.)^ the so-
called pharynx of Weismann and Van Rees, described by them in
the muscidian larva.

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266 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

The cephalic imaginal discs now grow rapidly, and soon attain
the form which characterizes them in the young larva. The median,
ventral disc has begun to invaginate (PL 6, Fig. 34, dsc, ce. m.), and
a cross-section through this region (PL 6, Figs. 36 and 37, dsc, ce, m,)
shows that the invagination is paired. (Compare these cross- sections
with Figure 34, where the positions of the cross-sections shown in
Figures 36-44 are indicated by the corresponding numbers, 36-44.)
The later history of this* disc (see Pratt, '93) shows that these paired
outgrowths of the ventral invagination develop during the larval
period into deep pockets, that a long projection springs from the
bottom of each of them, and that subsequently the pockets fuse in
the median plane and the projections develop into the proboscis of
the imago.

The muscular sucking tongue is now an internal organ (as may
be seen in Figures 34, 37 and 38), and projects free into the
pharynx. Figure 39 shows the base of the sucking tongue where
it is continuous on each side with the pharyngeal wall. The open
space dorsal to it is the forward end of the median portion of the
lumen of the dorsal discs (of, m.), where it communicates with the
pharynx. In the stage of development represented by Figure 30
(PL 5) , this opening (of, in.) leads to the outside of the animal's body.
The paired nervous masses below the oesophagus are portions of
the circum-oesophageal or cerebral commissures (Fig. 39, corns, ch) .
The section shown in Figure 40 (PL 7) also passes through the
base of the sucking tongue. The median portion of the fused dor-
sal discs (dsc, ce,) is much broader here, extending quite across the
embryonic body, while between it and the oesophagus {oe.) are
the muscle-mass of the tongue {lug,) and the cerebral nerve-
mass {(/n, cb.). The latter is in close contact with the ventral sur-
face of the disc, and with the tongue muscles quite encloses the
oesophagus. Figure 41 represents a section made immediately back
of the sucking tongue. The median portion of the fused dorsal
discs {dsc, ce,) is still shown, and between it and the oesophagus are
the median nerve {n, m,) — which joins the sucking tongue with the
. ganglia allata — and the two cerebral nerve-masses {(/n, cb,). Fig-
ure 42 shows a section considerably farther back (compare Fig. 34) ,
passing through the paired portions of the dorsal discs (dsc, ce.),
and also exhibiting the structures seen in the previous section. The
next figure (Fig. 43) shows the paired portions of the dorsal discs
{dsc. ce.) distinct from each other, though still in contact with the

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PR AIT: IMAGINAL DISCS. 267

cerebral nerve-masses {gn, cb.), as is also shown in the parasagittal
section (PI. 5, Fig. 33) . The ganglia allata (^n. ala,) and their
fusion to form the median nerve are shown in Figure 43. The
forward end of the heart (cr,) is also cut. Figure 44 represents a
section posterior to the dorsal discs and through the hinder part of
the ganglia allata. Mesoderm does not tnake its appearance in
the cephalic discs, nor do nerves or tracheae enter them.

The process of the involution of the embryonic head of Melopha-
gusj by which the ventral cephalic disc, the sucking tongue, and
the median opening of the dorsal cephalic discs, are changed from
external to internal organs, and the head of the animal telescoped
into the thorax, is really the final act of a longer operation. The
head of the embryo, which, in my opinion, is represented by the
head-fold of the germinal band, begins to disappear when the stomo-
deum begins its migration towards the anterior end of the embryo.
The involution of the head has been observed in only one other
representative of the brachycerous Diptera besides Melophagus,
viz. in Musca vomitoria by Weismann ('64) . Here the disappear-
ance of the head is also a gradual process. The embryonic head
gi*adually grows shorter and at the same time loses its segmenta-
tion, its posterior edge moves forward until finally its anterior
portion invaginates into the mouth, forming the so-called larval
pharynx. The dorsal cephalic discs then appear as appendages of
the dorsal pharyngeal wall. But Weismann did not observe the
method by which these structures originate.

2. 27ie Thoracic Discs. — Six pairs of thoracic imaginal discs
make their appearance in the embryo of Melophagus. Three pairs*
are dorsal and three pairs ventral, a dorsal and a ventral pair
belonging to each of the pro-, meso-, and metathoracic segments,
respectively. The three ventral pairs give rise during metamor-
phosis to the three pairs of imaginal legs ; the dorsal metathoracic
pair to the rudiments of the balancers, and the dorsal meso- and
pro thoracic pairs to no adult structiu^es, they being rudimentary
structures. All of these discs first appear late in the embryonic life
of the insect; namely, at about the time of the involution of the head,
— at a time when the foimation of the dorsal cephalic discs is prac-
tically complete, when the back of the embryo is closed, and the
wall of the intestinal tract is formed.

The earliest appearance of the thoracic discs figured is that
shown for the embryo represented in Figure 32 (PI. 5), in which the

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2r)8 PROCEEDINGS: BOSTON SOCIETY NATUR.\X HISTORY.

involution of the head is in progress. Three pairs of thickenings
of nearly equal size appear in the ventral ectoderm of the forward
end of this embryo. In the next stage shown (PL 6, Fig. 35), in
which the involution of the head is completed, these thickenings
have begun to invaginate, and three additional pairs of thickenings
have made their appearance in the dorsal ectoderm at the forward
end of the embryo. Only the dorsal j)rothoracic tliickening ap-
pears in Figure 35 ; it lies immediately in front of the dotted line
leading from the letters dsc, ce. The invagination of each ventral
oisc begins at its posterior border, as shown in the ventral meta-
thoracic disc in Figure 35 ; then the anterior border sinks in, as
shown in the mesothoracic disc in this figure ; finally the entire disc
sinks beneath the surface, as is shown in the prothoracic disc in the
same figure. As this figure clearly indicates, the invagination of
the three discs is not simultaneous, but the more anterior the disc
the earlier the invagination. Complete invagination rapidly follows
(PI. 7, Fig. 45) ; the disc at once separates itself from the ecto-
derm, and the opening made in the ectoderm by the invagination
closes (PL 7, Fig. 46) . It is while the discs are in this condition
that the embryo leaves the egg-envelopes, and the discs remain in
this condition during the entire larval life of the insect.

The dorsal thoracic discs do not invaginate during the embryonic
life of the animal. The meso- and metathoracic discs invaginate
in the young larva, and become detached from the ectoderm, as
do the ventral discs (PL 7, Fig. 40). The prothoracic discs do not
advance beyond the stage of development represented by figure 45,
but remain thick-walled pockets of the ectoderm. They are also
lined with a cuticula, continuous with that of the rest of the larval
ectoderm, and the different moultings take place from the pockets
as from the rest of the insect. In my preliminary paper (Pratt,
'97) I have stated that the dorsal prothoracic discs have a larval
origin, but later examination of the material has shown this to be
an error. As is the case of the cephalic discs, mesoderm does not
appear in the thoracic discs during the embryonic life of the insect,
nor do nerves or tracheae enter them.

3. The Discs of the External Genitalia. — These discs first
appear in the embryo shortly before it leaves the egg-membranes.
Two pairs of ectodermal thickenings then appear immediately
in front of the anus, — a larger posterior and a smaller anterior
pair. The former lie just in front of, and partly embracing,

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PRATT: IMAGINAL DISCS. 269

the anus and rectum (PL 2, Fig. 48, dsc, gen, p,) , and the latter
inimediately in front of it.

These two pairs of discs do not make their appearance simul-
taneously, the larger, posterior pair being the first to appear. These
have the form of two invaginations, one on each side of the anus
(Fig. 48), which project forward and fuse on the ventral (an-
terior) side of the end-intestine, forming a single flattened sac,
which partly encircles it (Fig. 49, dsc, gen, p,) , At its anterior
end this sac separates into two parts, being in this region again
paired (Fig. 50, dec, gn, p.) . It is in the outer ectoderm at this point
that the anterior discs finally make their appearance in the form
of a pair of ectodermal thickenings (Fig. 50, dec. gn. a,) , These
do not, however, develop further in the embryo ; but in the larva
they assume a sac-like form and are detached from the ectoderm.
A description of the development of these discs in the larva has
already been published (Pratt, '93) .

4. The Discs of the Internal Organs, — The imaginal discs of
the internal organs and of the abdominal hypodermis do not appear
during the embryonic life of the insect.

Methods.

The greater part of this investigation was carried on in the
Zoological Laboratory of Harvard University under the direction of
Prof. E. L. Mark. It was completed at Haverford College. Most
of the material was obtained in Germany, but a portion of it came
from the vicinity of Cambridge, Mass:, and of Haverford, Pa.

The eggs were obtained by dissection from the maternal uterus,
in which they develop. It was found to be impossible to remove
them in the fresh condition without injury, on account of the
extreme delicacy of the chorion. Consequently, in each case the
mother insect was killed by decapitation ; the abdomen was slit
open and then plunged into a warm corrosive-sublimate solution.
This fixed the %g^y and at the same time hardened it so that it
could be removed from the uterus. Most of the staining was done
with borax carmine, which was found sufficient for all ordinary
purposes, but Mayer's acid carmine and Ehrlich's haematoxylin
also were used.

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270 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

BIBLIOGRAPHY.

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'91. Recherches sur le developpement postembryonnaire, I'anatomie et les
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'32-'47. Handbuch der Entomologie. Berlin.
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Gonin, J.
'94. Recherches sur la metamorphose des l^pidopt^res. Bull. soc. vau-
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Graber, V.

'77-'79. Die Insekten. Zweiter Theil. Mttnchen.

'89. Vergleichende Studien tiber die Embryologie der Insecten und besond-
ers der Musciden. Denkschr. k. Akad. Wiss. Wien. math.-naturw. CI.,
Bd. 66, pp. 267-314. Taf. 1-10.
Grassi, B.

'84. Intomo alio sviluppo delle api nelP uovo. Atti accad. gloenia scL nat.
Catania, vol. 18, pp. 146-222, Tav. 1-10.

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PRATT: IMAGINAL DISCS. 271

Heider, K.
'89. Die Embryonalentwicklung von Hydrophilus piceus L. Jena. 98 pp.,
13Taf.
Henking, H.
'88. Die ersten Entwicklungsvorgftnge im Fliegenei und freie Kernbild-
ung. Zeitsclir. f. wiss. Zool., Bd. 46, pp. 289-336, Taf. 23-26.
Herold, M.
'15. Entwicklungsgeschiclite der Schmetterllnge anatomisch und physiolo
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Heymons, R.
*95. Die Embryonaleutwickelung von Dermapteren und Orthopteren, etc.

Jena. 136 pp., 12 Taf.
'97. Ueber die Zusammeusetzung der Insecteu-Kopf. Sitzungsber. Ges.
naturf. Fr. Berlin, no. 7, pp. 119-123.
Korotneff, A.

'85. Die Embryologie der Gryllotalpa, Zeitschr. f. wiss. Zool., Bd. 41, pp.
670-604, Taf. 29-31.
Korschelt, E., & Heider, K.
'92. Lehrbuch der vergleichenden Entwicklungsgeschichte der wirbellosen
Thiere. Jena*.
Kowalevsky, A.
'85. Beitrftge zur nachembryonaleu Entwicklung der Musciden. Zool.

Anz., Jahrg. 8, pp. 98-103, 123-128, 153-157.
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Ktinckel d' Ilerculais, J.

'75. Recherches sur 1' organisation et le dfiveloppement des volucelles.
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Landois, H.

'71. Beitrftge zur Entwicklungsgeschichte der SchmetterlingsflUgel in der
Raupe und Puppe. Zeitschr. f. wiss. Zool., Bd. 21, pp. 306-324, Taf. 23.
L6caillon, A.

'98. Recherches sur I'oeuf et sur le dfeveloppenient embryonnaire de
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Leuckart, R.

'58. Die Fortpflanzung und Entwicklung der Pupiparen. Abh. der naturf.
Ges. iu Halle, Bd. 4, pp. 1-81, Taf. 1-3.
Lyonnet, P.

1760. Traits anatomique de la chenille qui ronge de saule. La Haye.
Metschinkoff, E.
*83. Untersuchungen tiber die intracellulftre Verdauung bei wirbellosen
Thieren. Arb. zool. Inst. Wien, Bd. 6, Heft 2. pp. 141-168, Taf. 13, 14.
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'92. Der Rtissel der Diptera pupipara. Arch. f. Naturg., Jahrg. 58,
Bd. 1, pp. 287-332, Taf. 15, 16.

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272 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

Pancritius, P.
'84. Beitrftge zur Kenntniss der FltiKelentwickelung bei den Insecten.
Inaug.-Dissertation. Konigsberg. 37 pp., 2 Taf.
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Bd. l,pp. 151-200, Taf. 6.
'97. ImiiginaJ Discs in Insects. Psyche, vol. 8, pp. 15-30.
Rees, J. van.
'84. Over intra-cellulaire spijsverteering en over de beteekenis der witte
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Swammerdam, J.

1737-38. Bybel der natuure, etc. Leydae.
Verson, E.
'90. Der SchmetterljnKsfltif^el und die sogeuannte Imaginalscheibe der-
selben. Zool. Anz., Jahrg. 13, pp. 116-117.
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Voeltzkow, P. A.
'89. Entwicklung im Ei von Musca vomitoria. Arb. zool.-zoot. Inst.
WUrzburg, Bd. 9, pp. 1-48, Taf. 1-5.
Wahl, B.

'99. Ueber das Tracheen-system und die Imaginalscheiben der Larve von
Eristalis tenax L. Arb. zool. Inst. Wien, Bd. 12, Heft 1, pp. 46-98. Taf.
4-8.
Weisraann, A.
'63. Ueber die Entstehung des vollendeten Insectes in Larve und Puppe.

Abh. der senckenb. natiirf. Ges., Bd. 4, pp. 227-260, Taf. 10-12.
'64. Die nachembryonale Entwicklung der Musciden nach Beobachtungen
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Beitrfi,ge zur Anatomic und Embryologie als Festgabe Jacob Henle, zum
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Taf. 10-12.
Wheeler, W. M.
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Joiirn. morph., vol. 3, pp. 291-374, pis. 15-21.
Witlaczil, E.
'84. Entwicklungsgeschichte der Aphiden. Zeitschr. f . wiss. Zool., Bd. 40,
pp. 559-696, Taf. 28-34.

Printed June, 1900.

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Pratt.— Imaginal discs.

EXPLANATION OF PLATES.

All figures are from preparations of Melophagus ovinus L., and were made
with the aid of a camera lucida. The mesoderm and organs derived from it
are colored blue, the yolk yellow.

Abbreviations.

anterior.

Vvag. tr.

tracheal invagination.

am.

amniotic rudiment.

la. g.

germinal plate.

an.

anus.

In0.

sucking tongue.

bVdrm.

blastoderm.

mr'py.

micropyle.

chn.

chorion.

ms^drm.

mesoderm.

coel.

body-cavity.

n. m.

median nerve.

corns, cb.

cerebral commissures.

n. V.

ventral nerve chord.

cr.

heart.

nl.

segmentation nuclei.

eras. stmd.

stomodeal thickening.

nl. vt.

yolk nuclei.

crs. ma'drm.

mesodermic ridge.

oe.

oesophagus.

crs. m-v.

mid-ventral ectodermic

of. m.

median opening of paired

ridge.

cephalic disc.

dorsal.

or.

mouth.

dsc. al.

wing disc.

P-

posterior.

(Uc. ce.

paired cephalic disc.

phy.

pharynx.

dsc. ce. m.

median cephalic disc.

pli. ca.

taU-fold.

dsc. gen. a.

anterior genital disc.

pli. ce.

head-fold.

dsc. gen. p.

posterior genital disc.

prd.

proctodeum.

dsc. pd. 1,

prothoracic leg disc.

stmd.

stomodeum.

dsc. pd. 3.

mesothoracic leg disc.

St. pr'pl. pi.

peripheral protoplasmic

dsc. pd. 3.

metathoracic leg disc.

layer.

en'drm.

endoderm.

sul. m-v.

mid-ventral groove.

gl. sal.

salivary glands.

tae. rns^drm.

mesodermic bands.

gn. dla.

ganglia allata.

tb. mpg.

Malpighian tubules.

gn. cb.

cerebral ganglia.

tr.

tVachea.

gn.f.

ganglion froutale.

trt. pr^pl.

protoplasmic paths.

gran. vt.

yolk granules.

ventral.

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Pbatt. — Imag^inal discs.

PLATE 1.

Fig. 1. Longitudinal section of a segmenting egg, in a plane nearly coinciding

with the sagittal plane of the embryo. X 78.
Fig. 2. Similar section of a somewhat older segmenting egg. X 78.
Fig. 3. A portion of the section shown in the preceding figure, seen under a

higher magnification. X 760.
Fig. 4. Sagittal section of a segmenting egg in which all parts of the peripheral

layer of protoplasm are occupied by nuclei. X 78.
Fig. 6. A portion of the preceding section seen under a higher magnification,

many of the nuclei in process of division ; the peripheral layer of

protoplasm with shallow grooves surrounding the nuclei. X 760.
Fig. 6. A portion of a section of a segmenting egg showing the deepening of

the grooves in the peripheral layer of protoplasm to form the cells

of the developing blastoderm. X 760.
Fig. 7. A portion of a section showing the completely formed blasttxierm.

X876.

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Plate 1.

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PLATE 2.

Fig. 8. Cross-section of an embryo showing the origin of the mesoderm.

X130.
Fig. 9. Sagittal section of an embryo showing the formation of the head- and

tail-folds. X 100.
Fig. 10. Sagittal section of an embryo showing formation of proctodeum.

XIOO.
Fig. 11. Dorsal aspect of an embryo in which both proctodeum and stomodeum

have been formed. X 100.
Fig. 12. Semi-diagrammatic sagittal section of an embryo in which both proc-
todeum and stomodeum have been formed. X 100.
Fig. 13. Surface view, lateral aspect, of an embryo of the same stage as that

of Fig. 12. X 100.
Fig. 14. Cross-section through the stomodeum of an embryo of about the same

age as that shown in Figs. 12 and 13. X 130.
Fig. 15. Cross-section through the middle of the same embryo as the one shown

in the preceding figure. X 130.
Fig. 16. Cross-section through the proctodeum of the same embryo as that

shown in Fig. 14. X 130.
Fig. 48. Cross-section of an old embryo, passing through the anus and showing

the invaginations of the posterior genital discs. X 220.
¥ig. 49. Cross-section passing through the end-intestine and middle portion of

the genital discs. From the same en^bryo. X 220.
Fig. 60. Cross-section passing through the anterior genital discs of the same

embryo. X 220.

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PLATE 8.

Fig. 17. Surface view, dorsal aspect, of the head-fold of an embryo, showing

the first appearance of the cephalic imaginal discs, x 100.
Fig. 18. Cross-section through the stomodeum of an embryo somewhat older

than that represented in Fig. 17. X 130.
Fig. 19. Cross-section of the same embryo as that shown In Fig. 18, taken at

some distance further back, showing the cephalic discs where they are

deepest. X 130.
Fig. 20. Cross-section through the middle of the same embryo. X 130.
Fig. 21. Cross-section through the proctodeum of the same embryo. X 130.
Fig. 22. A somewhat oblique, but nearly sagittal, section of an embryo, the

stomodeum of which has reached a terminal poflition. X 130.

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Pratt- I MAG] NAL Disc:

Plate 3.

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PLATE 4.

Fig. 23. Parasagittal section of the same embryo as that shown in the preced-
ing figure. X 130.

Fig. 24. Cross-section through the extreme anterior end of an embryo some-
what older than that shown in the two preceding figures. X 200.

Fig. 25. Cross-section through the base of the sucking tongue of the same
embryo. X 200.

Fig. 26. Cross-section through the cephalic discs of the same embryo. X 200.

Fig. 27. Surface view of a wax model representing the anterior end of an
embryo of about tlie same age as that shown in Figs. 22 and 23. X 300.

Fig. 28. Surface view, lateral aspect, of the head end of an embryo of the
same age as that shown In Figs. 22 and 23. X 200.

Fig. 29. A slightly oblique sagittal section of an embryo, older than that
shown in Fig. 22, in which the involution of the head has begun.
X 130.

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PLATE 6.

Fig. 30. Anterior portion of the section represented in Fig. 29, seen under a

higher magnification. X 200.
Figs. 31 and 32. Parasagittal sections of the same embryo, Fig. 32 being

from the more lateral section. X 200.
Fig. 33. Parasa^ttal section of the same embryo as that shown in Figs. 34 and

36. It is more lateral in position than Fig. 36. X 200.

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PLATE 6.

Fig. 34. A slightly oblique sagittal section of an embryo in which the Involu-
tion of the head lb completed. X 200.

Fig. 36. Parasagittal section intermediate in position between those of Figs.
83 and 34 and from the same embryo as those figures. X 200.

Figs. 36-39. Cross-sections of the forward end of an embryo of about the same
age as that shown in the preceding figures. The region of the embryo
from which each of the sections is taken is indicated in Fig. 34 by
the number of the figure representing the section. X 200.

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PLATE 7.

Tigs. 40-44. Cross-sections of the forward end of the same embryo as that
shown in the preceding four figures. The region of each section is
indicated in Fig. 34, PI. 7, by the number of the figure which repre-
sents the section, x 200.

Fig. 45. Frontal section passing through an invaginating thoracic disc. X 340.

Fig. 46. Similar section of a completed thoracic disc. X 340.

[There is no Figure 47. Figures 48-60 are on Plate 2.]

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Proceedings of the Boston Society of Natural EUstorj.
Vol. 29, No. 14,
pp. 273-322.

GLACIAL EROSION IN FRANCE, SWITZERLAND AND NORWAY.

By William Morris Davis.

WiTHM

THREK PLATKS.

BOSTON:
PRINTED FOR THE SOCIETY.

July, 1900.

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NO. 14. — GLACIAL EROSION IN FRANCE, SWITZER-
LAND AND NORWAY.

By William Morris Davis.

' Introductory, — Eighteen years ago I presented to this Society
an essay on Glacial Erosion, in which my own observations were
supplemented by a review of all that I could find written on the
subject, in the hope of reaching some safe conclusion regarding
what was then (as it is still) a mooted question. Although recof^-
nizing effective erosion to depths of "a moderate number of feet"
where ice pressure was great and motion was rapid, in contrast to .
deposition where pressure and motion were reduced and where the
amount of subglacial drift was excessive, I could not at that time
find evidence to warrant the acceptance of great glacial erosion,
such as was advocated by those who ascribed Alpine lakes and
Norwegian fiords to this agency. In a retrospect from the present
time, it seems a^ if one of the causes that led to my conservative

.position' were the extreme exaggeration of some glacialists, who
found in glacial erosion a destructive agency competent to accom-
plish any desired amount of denudation — an opinion from which
I recoiled too far. Since the publication of my pi-evious essay I
had gradually come to accept a greater and greater amount of

. glacial erosion in the regions of active ice motion ; but in spite of
this slow change of opinion, the maximum measure of destructive
work that, up to last year, seemed to me attributable to glaciers
was moderate ; and it was therefore with great surprise that I then
came upon certain facts in the Alps and in Norway which demanded
wholesale glacial erosion for their explanation. The desire of some
years past to revise and extend my former essay then came to be a
duty, which it is the object of this paper to fulfil.

My former revision of the problem divided the arguments for
glacial erosion under four headings: observations on existing
glaciers and inferences from these observations; the amount and
arrangement of glacial drift; the topography of glaciated regions;
and the so-called argument from necessity, — that is, the belief
that glaciers must have done this and that because nothing else

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274 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

competent to the task could be found. It is not possible for me at
present to review all the new material pertinent to the whole
problem; attention can be given here chiefly to a few examples
under the third heading. •

A Glaciated Valley in Central France, — It is evident that, if it
were possible to obtain a definite idea of the preglacial topography
of a glaciated district, the amount of glacial work might be readily
determined as the difference between the preglacial and the present
form ; independent evidence sufficing to prove that general denu-
dation of the rocky crust in the brief postglacial epoch had been
inconsiderable. This method leads one to conclude that in gen-
eral the topography of southern New England has not been
strongly modified by glacial action ; for we find here on the whole
the same maturely dissected upland that prevails in regions of simi-
lar structure outside of the glacial boundary; the uplands being
explained as parts of an uplifted peneplain of late Mesozoic date,
and the valleys as the work of ordinary erosion in a part of Terti-
ary time : but this method of measuring glacial erosion by dating
topographic forms had not been developed twenty years ago.
Strong glacial erosion may, however, be expected in New England
where ice motion was locally accelerated, as through the notches
of the White Mountains. Again, in the glaciated area of the Cen-
tral Plateau of France, I had opportunity in January, 1899, of
seeing a valley that had been locally modified to a determinate
amount by a glacier that once descended northwest from the
Cantal along the valley of the Rhue to the junction of the latter
with the Dordogne. Outside of the glaciated area, the valleys of
the plateau — an uplifted and sub-maturely dissected peneplain,
mostly of crystalline rocks — frequently follow incised meandering
courses, in which the steep concave slopes are regularly opposed to
the gentler convex slopes ; the latter being spur-like projections of
the uplands, advancing alternately from one and the other side of
the valley. Valleys of this kind are singularly systematic in form,
as the result of the combined downward and outward cutting by
their streams which, already winding or meandering when the
erosion of the valleys began, have increased the width of their
meander belt while they deepened their valleys. On entering tlie
glaciated valley of the Rhue, it is found that the regularly descend-
ing spurs of the non- glaciated valleys are represented by irregular
knobs and mounds, scoured on their up-stream and plucked on the
down-stream side ; and that the cliffs formed where the spurs are

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DAVIS: GLACIAL EROSION.

275

cut off, as in Fig. 1, are sometimes fully as strong as those which
naturally stand on the opposite side of the valley. The spurs
generally remain in sufficient strength to require the river to follow
its preglacial serpentine course around them, but they are some-
times so far destroyed as to allow the river to take a shorter course

Fig. 1. The glaciated valley of the Rhue.

through what was once the neck of a spur.^ The short course is
not for a moment to be confounded with the normal cut-offs through
the narrowed necks of spurs, such as are so finely exhibited in the
meandering valleys of the Meuse and the Moselle. The short
courses are distinctly abnormal features, like the rugged knobs to
which the once smooth- sloping spurs are now reduced.

It was thus possible in the valley of the Rhue to make a definite
restoration of preglacial form, and to measure the change produced
by glaciation. The change was of moderate amount, but it was
highly significant of glacial action, for it showed that while a slender,
fast-flowing stream of water might contentedly follow a serpentine
course at the bottom of a meandering valley, the clumsy, slow-
moving stream of ice could not easily adapt itself to so tortuous a
path. The more or less complete obliteration of the spui-s was the
result of the effort of the ice stream to prepare for itself a smooth-
sided trough of slight curvature ; and if the rocks had been weaker,
or if the ice had been heavier, or if the glacial period of the Cantal
had lasted longer, this effort might have been so successful as to
have destroyed all traces of the spurs. Fortunately the change
actually produced, only modified the spurs, but did not entirely

1 The short-cuts are sometimes narrow gorges incised in the half-consumed spurs ; and
in such cases, tlie displacement of the Rhue from its former roundabout course is
probably to be exi)Iained by constraint or obstruction by ice.

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276 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

destroy them ; and their rugged remnants are highly significant of
what a glacier can do.

Rocky Knobs in Glaciated Areas, — On thus generalizing the
lesson of the Rhue, it is seen that just before the complete oblitera-
tion of the spurs some of their remnant knobs may be isolated from
the uplands whence these preglacial spurs descended. It is out of
the question to regard the ruggedness of such knobs as an indication
of small change from their preglacial form, as has been done by
some observers. The ruggedness is really an indication of the
manner in which a glacier reduces a larger mass to smaller dimen-
sions, by plucking on the down-stream side as well as by scouring
on the up-stream side. It is possible that knobs in other glaciated
valleys than that of the Rhue may be of this origin ; they should
then be regarded not as standing almost unchanged and testifying
to the incapacity of glacial erosion, but as surviving remnants of
much larger masses, standing, hke raonadnocks above a peneplain,
as monuments to the departed greater forms. The two knobs at
Sion (Sitten) and the Maladeires, all detached from Mont d'Orge
in the upper valley of the Rhone, the hills of Bellinzona in
the valley of the Ticino, the rocks of Salzburg where the Sal-
zach emerges from the Alps, and even the Borromeo islands in
Lake Maggiore, may perhaps be thus interpreted. Rugged as these
knobs may be on the down-stream side, it would be an unreasonable
contradiction of the conclusions based on observations of many
kinds to maintain that their ruggedness was of preglacial origin.

The ice stream from the Cantal at one time expanded sufficiently
to flood the uplands bordering the valley of the Rhue,^ where it
produced changes of a most significant kind. The neighboring
unglaciated uplands are of systematic form ; broad, smoothly arched
masses rise, round-shouldered, between the narrow valleys that are
incised beneath them ; the uplands are as a rule deeply soil-covered,
and bare ledges prevail only on the stronger slopes of the young
valleys that have been eroded since the peneplain was raised to its
present upland estate. But within the glaciated area near the
Rhue, the broadly rounded forms of the uplands are replaced by a
succession of most irregular rocky knobs, from which the preglacial
soils have been well scom'ed away, as in Fig. 2. This seems to be
a form most appropriate to glacial action on a surface that had been

1 According to Boule ('96), the glaciation of the uplands and of the valleys was sepa-
rated by an interplacial epoch, but I did not have occasion to inquire particularly inta
this aspect of the problem

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277

weathered to variable depths in preglacial time. The ice action
sufficed to rasp away the greater part of the weathered material
and to grind down somewhat the underlying rock, often giving the
knobs a rounded profile; but it did not nearly suffice to reduce the

Fig. 2. Glaciated knobs on the Central Plateau of France.

rocky surface to an even grade. The ice action seems here to have
resembled that of a torrent which might sweep away the waste on
a flood plain and lay bare and erode the rock ledges beneath ; but
whose duration was not sufficient to develop a graded floor appro-
priate to its current.

Another example of this kind seems to occur where the huge
glacier of the Inn, escaping from its well enclosed channel within
the mountains, once spread forward in a great fan of ice over the
foot-hills at the northern border of the Alps and crept out upon
the piedmont plain. The glance that I had at this foot-hill district
from a passing train gave me the impression that its ruggedness was
much greater than usually obtains along the mountain flanks ; as if
the rolling hills of preglacial time had been scoured to an increas-
ing roughness by an overwhelming ice-flood that would, if a longer
time of action had been permitted to it, have worn down all the
inequalities to a smooth, maturely graded floor.

The Vcdley of the Ticino, — My first entrance into the Alps last
year was from the south by the valley of the Ticino. Thirty-one
years before I had followed the same valley and admired its bold
sides and its numerous waterfalls; but at that time nothing was
noticed that seemed inappropriate to the general idea of the erosion

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278 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

of valleys by their rivers. Thirty years is a long enough time for
one to learn something new even about valleys, and on my second
visit it was fairly startling to find that^the lateral valleys opened on
the walls of the main valley of the Ticino five hundred feet or more
above its floor, and that the side streams cascaded down the steep
main-valley walls in which they have worn nothing more than nar-
row clefts of small depth. This set me wondering, not only as to
the meaning of so peculiar an arrangement of valleys and streams,
but also as to the reason why so peculiar an arrangement should
not have sooner attracted attention as an exceptional characteristic
of Alpine topography. Playfair long ago, when describing the
relation of side valleys to their trunk, showed clearly that they had

" such a nice adjustment
of their declivities that
none of them join the
principal valley either on
too high or too low a
level : a circumstance
which would be infinitely
improbable if each of
these vallies were not the
work of the stream that
flows in it" ('02, 102);
yet the whole course of
the passing century has
hardly sufficed to make
full application of this
law. So much latitude
is usually allowed in the
relation of branch and
trunk valleys that hun-
dreds of observers, many
of whom must have been
cognizant of Playf air's
law, have made no note
of the extraordinary exceptions to it that prevail in the glaciated
valleys of the Alps. Even the most pronounced advocates
of glacial erosion, with a few exceptions to be noted below,
have been silent regarding the remarkable failure of adjustment
between the declivities of lateral and main glaciated valleys.
Indeed, in reviewing the writings of those who have accepted a

''^^2^£^^' "I

Fii?. 3.

Val d'Osogiia, a hanging lateral valley
of the Ticino.

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DAVIS: GLACIAL EROSION. 279

large measure of glacial erosion, one must be struck with the undue
attention that they have given to lake basins and the relative inat-
tention to valleys. This disproportion is probably to be explained
as a result of the greater contrast that prevails between a river and*
a lake than between a river and its branch ; it is perhaps for this
reason that the attention of geologists and geographei-s has gener-
ally been directed to the origin of lakes rather than to the relation
of branch and trunk streams, even when the former cascade from
their lateral valleys into the main valley. That glacial erosionists
made so little claim for the general deepening of glaciated valleys
while they demanded a great deepening of those parts of valleys
which have been scoured down to form lake basins, has always
seemed to me a difficulty in the way of accepting the demanded
measure of lake-basin erosion ; and this difficulty was supported by
the well-attested observation that the side slopes of glaciated valleys
manifest no marked or persistent increase of declivity in passing
from above to below the limit of glaciation. If glaciers had scoured

out deep lake basins, like
those of Maggiore and

^ ^^/nmh Geneva, they ought to have

significantly deepened the
valleys up-stream from the
lakes; and if the valleys were
thus significantly deepened,
it seemed as if their slopes
Fig. 4. Section of a glaciated valley. ^\iOM\^ be steeper below than

above the limit of glacial
action. The denial of the latter requisite seemed to me to carry
with it the denial of the two preceding suppositions.

Features of Strongly Glaciated Valleys, — It is true that the
uppermost limit of glaciation, Q R, Fig. 4, in Alpine valleys is
not attended by a persistent change in the steepness of the valley
sides, A E, C J ; but on descending well within the glaciated val-
ley, a very strong change may usually be found in the slope of
the valley walls. The larger valleys, once occupied by heavy
glaciers from the lofty central snow fields, are chai-acterized by
" basal cliffs,*' E F, J II, that rise several hundred or even a thou-
sand feet above their broad floors, and thus enclose what may be
called a " bottom trough,*' E F II J, half a mile or a mile wide.
The bottom trough of the Ticino, as seen when one looks up
stream towards Giornico, is shown in Plate 1, Figure A, The

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280 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

basal cliffs are comparatively straight- walled ; they have no sharp
spurs advancing into the valley floor. The rock floor, G, Fig. 4,
is buried by gravels, F H, to an unknown depth. It is only from
the benches above the basal cliffs that the valley sides flare open
with maturely inclined slopes; and it is at a moderate depth
beneath the level of the benches at the top of these basal cliffs
that the lateral valleys, D K, open on the walls of the main valley.

The bottom trough within the basal cliffs and beneath the lateral
valleys seems to be of glacial origin. It is in the first place a
characteristic feature of all the larger glaciated Alpine valleys, as
I am assured by Professors Penck, Brttckner and Kichter, with
whom the matter was carefully discussed last summer. The non-
glaciated valleys manifest no such peculiar form. It is not simply
that the terminal portion, J B K, of a lateral valley has been cut
off by the glacial widening of the paain valley floor; the main
valley has been strongly deepened, as is assured by the relation of
its floor, F H, to the prolongation of the floor of the lateral valley,
KB. The first may be several hundred feet — indeed in some
valleys, a good thousand feet — below the second. The lateral
valleys must have once entered the main valley at grade, for the
flaring sides of the main valley indicate maturity ; the side slopes,
AE, CJ, must have once met at B. Even the lateral valleys
have an open V section, proving that their streams had cut down
to a graded slope, D B, that must have led them to an accordant
junction with the main river. Nothing seems so competent as
glacial erosion to explain the strong discordance of the existing
valleys.

The lateral as well as the main valleys have been glaciated, but
the former do not exhibit changes of form so distinctly as the
latter: in the Ticino system the lateral valleys did not, as far as I
saw them, seem to have been much affected by glaciation, a fact
that may be attributed to the small size of their branch glaciers in
contrast with the great volume of the trunk glacier. There is no
sufficient evidence that the valley floor between the basal cliffs has
been faulted down, after the fashion of a grahen ; for although this
origin is advocated by Rothpletz (*98, 237) for the Linththal, the
evidence that he adduces for the limiting faults is not agreed to
by Alpine geologists in general, and the persistent association of
the bottom troughs with the crooked course of pre-existent,
maturely open valleys involves special conditions of faulting that
cannot be accepted without the strongest evidence.

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DAVIS: GLACIAL EROSION. 281

It is not satisfactory to explain the bottom trough as having
been worn out by normal trunk-river erosion, leaving the side
streams as it were hanging or suspended above them, for to admit
such an origin would be to go counter to all that has been learned
regarding the systematic development of valleys. Here it is with
regret that I must differ from the opinion of two eminent Swiss
geologists who explain the deepening of the main valleys by a
revival in the erosive power of the rivers as a result of a regional
uplift, while they regard the hanging lateral valleys as not yet
accordantly deepened by their smaller streams. It is true that
narrow trenches are cut in the floors of the hanging valleys, show-
ing that their streams have made some response to the erosion of
the bottom though in the main valley, and if the bottom trough
were a narrow canyon, this relation of trunk and branch streams
might be considered normal; but if the breadth as well as the
depth of the bottom trough had been acquired by normal river
erosion, the side valleys should now, it seems to me, have been
trenched much deeper than they are, to some such slope as ST,
Fig. 4.

The opinions of Rtitimeyer and Heim on this question are as
follows : — Rtitimeyer gave an excellent account of hanging lateral
valleys thirty years ago in his description of the valley of the
Reuss (*69, 13-24). He recognized benches or ThaUtufen on each
side of the valley above the basal cliffs of the existing bottom
trough, and regarded them as the remnants of a former, wide open
valley floor. Side valleys of moderate fall enter the main valley
about at the level of the Thalstufen^ and their waters then cascade
down over the basal cliffs to the Reuss. Glacial erosion is dismissed
as incompetent to erode the bottom trough ; indeed, the time of
glacial occupation of the valley is considered a period of rest — a
sort of "pupa stage" — in its development. The discordance of
main and lateral valleys is ascribed entirely to the differential
erosion of their streams. Heim's views on this matter are to be
found in his ^^ Mechanismus der Gehirgshildung^^ ('78, 1, 282-301)
and in an article *' JJcher die Erosion im Gehiete der Reuss " ('79).
He recognizes that the bottom troughs have been excavated in the
floors of pre-existing valleys, whose stream lines had been reduced
to an even grade (profile of equilibrium, " GleichgercichtsUnie'')
and whose lateral slopes had been maturely opened. The side
streams must at that time have eroded their valleys deep enough
to enter the main valley at accordant grade as stated above. Since

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282 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

then, it is concluded that an elevation of the region has caused a
revival (Neubelebimg) of the main river ; and the present greater
depth of the main valley is, according to Heim, merely the natural
result of this revival, while the smaller side streams have not yet
been able to deepen their valleys. The height of the ThcUstuJen
or remnants of the former valley floor, seen in the benches above
the basal cliffs of the bottom trough, is taken as a measure of the
elevation that the mountain mass has suffered.

Apart from the improbability that the deepening of a bottom
trough by a revived main river could truncate so many lateral valleys
with so great nicety as is repeatedly the case, leaving their streams
to cascade down in clefts but slightly incised in the main valley
walls, the following considerations lead me to reject the possibility
of explaining the discordance between side and main streams by a
normal revival of river action.

Mekition of Trunk and Branch Valleys, — The general accord-
ance of maturely developed main and lateral valleys in non-glaciated
regions, as recognized by Playfair, is today fully established by
innumerable observations in many parts of the world. Truly,
during the attainment of mature development, it is possible that a
large river may outstrip a small branch stream in the work of deep-
ening its valley, but the discordance thus produced can prevail only
during early youth ; for as soon as the main river approaches grade
the further deeping of its valley is retarded, while at the same time
the steepened descent of the lateral streams at their entrance into
the main valley accelerates their erosive work. Hence, even if a
large trunk river has for a time eroded its valley to a significant
depth beneath the tributary valleys, this discordance cannot endure
long in the history of the river. Examples of such normal discord-
ance are to be found in non-glaciated regions only in the branch
streams of rivers that occupy very narrow canyons ; and even rivers
in canyons sometimes receive their branches at accordant grade, as
seems to be usually the case with the Colorado, if one may judge by
photogi-aphs. The narrow postglacial gorges cut by active streams,
habitually receive their branches — when they receive any — from
hanging side gorges ; and an excellent example has long since been
on record in the gorge of Cattaraugus Creek in wegtem New York,
where a branch, the Canaserowlie, falls into the main gorge from a
side gorge of much less depth. Referring to this. Hall wrote: —
" In the more recently excavated channels we find the streams fall-
ing over the very edge of the cliff, having produced no perceptible

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DAVIS: GLACIAL EROSION. 283

recession in the margin of the fall" (*43,380). But however ap-
propriate a discordance of branch and trunk may be in early youth,
it cannot endure long enough to be associated with maturely opened
main valleys. It should be noted that discordance of side and main
valleys may also be found where a large river has lately been turned
to a new path, as in the normal progress of the capture of the upper
course of one river by the headward gnawing of a branch of another
river (see reference to Russell below) , or in the new arrangements
of drainage lines in a region from which a glacial sheet has lately
withdrawn. Furthermore, the valleys of very small wet-weather
streams are frequently discordant with the valley of a serpentine
river, if they enter it from the upland that is under-cut by the con-
cave bank of the river. But these cases cannot find application in
the hanging valleys of the Alps. The hanging valleys that open on
sea cliffs, such as those of Normandy, are of course quite another
matter.

Overdeepened Main Valley a and Hanging Lateral Valleys, —
Now it is characteristic of the bottom troughs of the glaciated
Alpine valleys that they are broad-floored ; they cannot be described
as canyons in any proper sense of that word : the walls are steep
enough, but they are too far apart. If the existing breadth of the
troughs had been acquired in the ordinary manner by the lateral
swinging of the main stream and by the lateral weathering of the
walls, the long time required for such a change would have amply
sufficed for the lateral streams to cut down their valleys to grade
with the main valley ; and their persistent failure to do so indicates
the action of something else than normal river work in the widening
of the main valley. This is the very kernel of the problem.

If a main valley were excavated along a belt of weak rocks, the
side valley might stand for some time at a considerable height above
the main valley floor. Certain hanging vaUeys in the Alps seem at
first sight to belong to this class, but such is not really the case.
For example, where the Linth flows into the Wallen See, the well-
defined bottom troughs of the river and of the lake both pass
obliquely through a syncline of strong lower Cretaceous limestone,
which forms cliffs on their walls. Side streams drain the high
synclinal areas; one such stream cascades from the west into the
Linth trough back of the village of Nafels ; another cascades from
the north into the Wallen See near its western end. The first
explanation for such falls is that they are normally held up on the
resistant limestone ; but it should be noted that the bottom troughs

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284 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

of the Linth and the Wallen See have been cut down and broadly
opened in the same limestones. If the troughs were of normal
river origin, the side streams also should have by this timB trenched
the limestones deeply, instead of falling over the limestone cliiTs at
the very side of the larger troughs. In the Ticino valley where
the side streams are most discordant, massive gneisses prevail ; the
structure is so nearly uniform over large areas that it affords no
explanation of the strong discordance between side and main
valleys.

It thus seems obligatory to conclude that the bottom troughs of
the larger Alpine valleys were deepened and widened by ice action.
This belief is permitted by the abundant signs of glacial erosion on
the spurless basal cliffs, and required by the persistent association
of over-deepened bottom troughs and discordant hanging lateral
valleys with regions of strong glaciation. The valley of the Ticino
manifests these peculiarities very distinctly, and I have recently
described them in some detail in a paper in Appalachia (1900) .

Subaerial Erosion during the Gldcial Period, — It should not
be imagined that the glacial erosion of troughs in valley floors was
necessarily so rapid that no significant subaerial erosion was accom-
plished during its progress. Ordinary weathering and down-hill
transportation of rock waste must have been in active operation on
the valley sides above the border of the ice-filled channels; and
the very fact that on the upper slopes of the mountains, pre-
glacial, glacial and postglacial erosion was similarly conditioned,
makes it difficult to distinguish the work done there in each of these
three chapters of time. In the diagrams accompanying this article
no indication of change from preglacial to postglacial outline on
the ui>per mountain slopes is indicated, because no satisfactory meas-
ure can be given to it.

Lake Lugano, — In the presence of a variety of evidence col-
lected for some years previous to my recent European trip, it had
been my feeling that the best explanation offered for the large lakes
that occupy certain valleys on the Italian slope of the Alps was
that they had resulted from what has been called valley-warping, as
set forth by Lyell, Heira and others. It was my desire to look es-
pecially at Lakes Maggiore, Lugano and Como with this hypothesis
in mind, and to subject it to a careful test by means of certain asso-
ciated changes that should expectedly occur on the slopes of the
neighboring mountains, as may be explained as follows.

On the supposition of moderate or small glacial erosion, a well-

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DAVIS: GLACIAL EROSION. 285

matured stage of dissection must have been attained in the district
of the Italian lakes in preglacial time ; for the main valleys are
widely opened, and even the lateral valleys have flaring slopes.
In a mature stage of dissection mountains should exhibit a well-
advanced grading of their slopes ; that is, their sides should be
worn back to a comparatively even declivity with little regard to
diversity of structure ; the descending streams of waste being thus
seen to correspond to the flood plains of graded rivers. The
agencies of weathering and transportation are delicately balanced
wherever graded slopes prevail ; and a slight tilting of the moun-
tain mass might suffice to disturb the adjustment between the sup-
ply and the removal of waste ; then all the steepened slopes would
soon be more or less completely stripped of their waste cover;
their rock ledges would be laid bare, although still preserving
the comparatively even declivity that had been gained under the
slowly moving waste.

If the lakes had been formed by warping, it is possible to deduce
with considerable accuracy the localities where the mountain slopes
would be steepened and stripped; namely, the northern slopes
about the southern end of the lakes, and the southern slopes about
the northern end ; but as far as I was able to examine the district
about Lake Lugano, no effects of such a warping and tilting were
to be detected. The submergence of lateral valleys about the
middle of the lakes is also, as has been well pointed out by Wallace,
a necessary consequence of the theory of warping; but although
the main valley floor is now deep under water, the side valleys
are not submerged. Failing to find evidence of warping, and
being much impressed with the evidence of deep glacial erosion as
indicated by the hanging lateral valleys of the overdeepened
Ticino, I examined the irregular troughs of Lake Lugano for similar
features, and found them in abundance.

One of the reasons why Lake Lugano had been selected for
special study was that it did not lie on the line of any master
valley leading from the central Alps to the piedmont plains ; hence,
if influenced by ice action at all, its basin must have been less
eroded than those of Como and Maggiore on the east and west.
But in spite of this peculiarity of position, Lugano received strong
ice streams from the great glaciers of the Como and Maggiore
troughs (see Glacial Distributaries, below), and its enclosing slopes
possess every sign of having been strongly scoured by ice action.
The sides of the lake trough are often steep and cliff-like for hun-

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286 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

dreds of feet above present water level, thus simulating the basal
cliffs of the Ticino valley ; while at greater heights the valley sides
lean back in relatively well-graded slopes, as in Plate 1, Figure B^
where the southern side of the northeastern arm of the lake, near
Porlezza, is shown. The angle at the change of slope is often well
defined, but it is independent of rock structure. Narrow ravines are
frequently incised in the basal cliffs, and alluvial fans of greater or
less size are built into the lake waters from the base of the ravines.

The northeastern arm of the lake, extending from the town
of Lugano to Porlezza, receives several cascading streams from
hanging valleys on its southern side, one of which is here shown in
Plate 2, Figure A, The side slopes of the hanging valleys are for the
most part flaring open and well graded, from which it must be con-
cluded that their streams had, under some condition no longer exist-
ing, ceased to deepen their valleys for a time long enough to allow the
valley sides to assume a mature expression ; and that since then the
bottom trough of the main arm of the lake has been eroded deep and
wide, with a very small accompanying change in the lateral valleys.
In other words, the side valleys were, in preglacial time, eroded to
a depth accordant with the floor of the master valley that tliey
joined, and since then the bottom trough has been eroded in the
floor of the master valley by a branch of the Como glacier. In
postglacial time the side streams have begun to trench their valley
floors, eroding little canyons ; but much of this sort of work must
be done before the side valleys are graded down even to the level
of the lake waters, much less to the level of the bottom of the lake.

The two southern arms of the lake lead to troughs whose floors
ascend southward to the moraines of the foot-hills, beyond which
stretch forward the abundant overwashed gravels of the great plain
of the Po.

I do not mean to imply that every detail of form about Lake
Lugano can find ready explanation by the mature glacial modifi-
cation of a mature preglacial valley system ; but a great number of
forms may be thus explained, and a beUef in strong glacial erosion
was forced upon me here as well as in the valley of the Ticino.
A detailed study of the Italian lakes with the intention of care-
fully sorting out all the glacial modifications of preglacial forms
would be most profitable.

Various Examples of Glaciated Valleys, — My excursions of
last summer showed me a number of over-deepened main valleys
and hanging lateral valleys in the Alps ; for example, those of the

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DAVIS: GLACIAL EROSION.

287

Fig. 6. True-scale cross-section
of the Lauterbrunnen valley.

Inn and of the Aar. Lakes Thun and Brienz receive numerous
cascades from hanging valleys that stand high above the water
surface. The valley of Lauterbrunnen also affords a conspicuous
illustration of a deep bottom trough enclosed by high basal cliffs
that rise to the edge of more open upper slopes ; the celebrated
Staubbach fall is the descent of a small lateral stream from its lofty
hanging valley (see extract from an article by Wallace, cited
below) , and the picturesque village of JVItlrren, M, Fig. 5, stands

on the flaring slope or Thalstufe of
the preglacial valley, just above the
great basal cliff of glacial origin. A
mile or so south of the village of
Lauterbrunnen, the Trummelbach,
T, Fig. 5, descends the precipitous
eastern wall from a hanging valley
whose floor is hundreds of feet above
that of the Ltltschine ; it is roughly
sketched in Fig. 6. Although the
lateral Trummelbach brings a large volume of water to the main
valley, it descends by a very narrow cleft in the rock face, a trifling
incision in the valley wall; while the main valley, whose trunk
stream did not seem to be more
than five times the volume of its
branch, is half a mile or more
broad, wide open and flat-floored.
The cross-section of the main
valley is over a thousand times as
large as that of the lateral cleft.
Such a disproportion of main
valley and lateral cleft is entirely
beyond explanation by the in-
equality of their streams ; and for
those who feel that they must
reject glacial erosion as the cause
of the disproportion, there seems
to be no refuge but in ascribing
the main valley to recent down-
faulting : a process that can hardly

be called on to follow systematically along the floors of the larger
glaciated valleys of the Alps, and to avoid the non-glaciated valleys
and the mountain ridges.

^ig. 0. Diagram of the gorge of
the Trummelbach, Lauterbrunnen
vallev.

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288 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

Certain well-known Alpine glaciers may be instanced as reaching
just beyond the end of a hanging lateral valley and thence cascading
into the deeper main valley. One is the Mer de Glace by Chamou-
nix ; its cascading end is known as the Glacier des Bois. Another
is the neighboiing Glacier des Bossons, from whose upper amphi-
theatre a steep tongue descends far below; like the waterfalls of
Norway, the tongue may be seen lying on the side slope from a
considerable distance up or down the main valley. A third example
is the Glacier of the Rhone, whose splendid terminal cascade is so
consjncuous from the road to the Furka pass. (These three I have
seen some years ago.) Possibly the Vernagt glacier is another of
the same kind ; its catastrophic overflows into the lower Rofen
valley have often been described. Doubtless many other examples
of this class might be named.

While engaged upon these observations in the Alps in the spring
of 1899, I sent a brief note about them to my esteemed friend, Mr.
G. K. Gilbert of Washington, telling him that all the lateral vallej's
seemed to be "hung up" above the floors of the trunk valleys.
His reply was long in coming to Europe, and, on arriving at last,
it was dated Sitka, Alaska, where Mr. Gilbert had gone as a mem-
ber of the Ilarriman Alaskan Expedition, and where my note had
been forwarded. He wrote that, for the fortnight previous to
hearing from me, he and his companions had been much impressed
with the discordant relations of lateral valleys over the waters of
the Alaskan fiords, and he suggested that such laterals should be
called "hanging valleys" — a term which I have since then adopted.
He fully agreed that hanging valleys presented unanswerable tes-
timony for strong glacial erosion, as will be stated in his forthcoming
report on the geology of the Expedition.

After leaving Switzerland, I had a brief view of the lake district
in northwest England, before crossing to Norway. The amount of
glacial erosion in the radiating valleys of the English lakes has
been much discussed, and as usual directly opposite views have
been expressed. Rugged rocky knobs were seen in abundance
about Ambleside and along the ridge separating the valley of
Thirlmere from St. John's Vale ; and the latter receives a hanging
valley from the east near Dalehead post-oflice. The famous falls
of Lodore seemed to descend from the mouth of a hanging valley
into Derwentwater. A model of the lake district, on exhibition at
Keswick, showed some other examples of lateral valleys that
seemed to stand above the floors of their main valleys, notably one

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DAVIS: GLACIAL EROSION. 289

coming from the south near the northeast end of UUeswater.
Since coming home I read the following in Marr*s " Scientific Study
of Scenery": — "We find in the Lake District a number of tribu-
tary valleys occurring in the hearts of the ridges, and opening out
far above the bottoms of the main valleys, discharging their waters
down the slopes in cascades. They are specially well marked on
the east side of Helvellyn, and a number of them also open into the
upper branches of Borrowdale." The explanation is that of Rtlti-
meyer and Heim : — " For a considerable period after the deepening
of the main valley, the minor valleys will end as definite gorges
some height above the floor of the main valley, and discharge their
waters in a series of cascades or falls down the side of the main
valley" (1900, 136).

One of my former students, Mr. W. B. Lloyd, has recently shown
me a number of photographs of the fiords of southern New Zea-
land, which he brought back from a visit to that distant country.
High cascades, plunging from hanging lateral valleys into the
broad waters of the fiords, are repeatedly shown ; the most striking
view is here reproduced in Plate 2, Figure £ showing Sterling Fall
leaping into Milford Sound.

Fiords and Hanging Valleys in Norway, — In Norway I had
the pleasure of making a ten days' cross-country excursion in com-
pany with Dr. Reusch, Director of the Norwegian Geological Survey.
We entered from Bergen through Hardanger fiord, and crossed the
highlands by the Ilaukelisaetr road to Skien on the southeastern
lowlands, thus making a general cross-section on which many char-
acteristic features were seen. Norway has long been known as a
land of waterfalls, but it is not generally stated with sufiicient clear-
ness or emphasis that many or most of the falls are formed by the
descent of streams from maturely opened trough-like hanging val-
leys which are abruptly cut off by the walls of the fiords. The
discordance between main and side streams is simply amazing. The
fiord valleys are frequently one or two miles wide ; the waters of the
fiords are of great depths, reaching 3000 feet in some cases. Even
when a side valley stands but little above sea-level, its floor may be
half a mile above the floor of the fiord. On passing inland beyond
the head of the fiord water, where the whole depth of the fiord
valley is visible, the side valleys may open more than a thousand
feet above the main valley floor. In many cases where the fiords
are enclosed by smooth walls, the cascading side streams have not
yet incised a cleft in the bare rock surface, so that their foaming

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290 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

waters are visible for many miles up and down the fiord. Streams
of considerable size sometimes plunge down from the rolling uplands
in whose edge they seem to have just begun to cut a cleft. Ab-
normal discordance of trunk and branch stream is, therefore, a
strongly marked characteristic of the Norwegian drainage. The
necessity for appealing to strong glacial erosion in explanation of
this prevailing discordance, may be set forth as follows.

Measure of Glacial Erosion in Norwegian Eiords, — The deep
valleys of Norway, partly occupied by sea water, are incised
beneath an uneven highland which bears so many hills and moun-
tains that it makes little approach to a peneplain, yet which here
and there shows so many broadly opened uplands between the bills
and mountains that it may be taken to represent the well-advanced
work of a former cycle of denudation when the region stood much
lower than it stands now. As a whole, a mature or late mature
stage seems to have been reached before a movement of uplift
introduced the present cycle. Let us now make two suppositions
regarding the work of normal river erosion in the preglacial part of
the present cycle, in order to determine, if possible, how much addi-
tional erosion must be attributed to ice in the production of existing
forms.

First, let it be supposed that the revived main rivers had incised
their valleys to the depth of the present fiords in preglacial time,
and that the discordance of main and side valleys now visible is the
appropriate result of the youth of the present cycle. If we recall
only the steepness of the fiord walls, this supposition might be justi-
fied, and thus the amount of glacial erosion needed to develop exist-
ing forms would be small. But it must not be forgotten that the
fiords, although often steep-walled, are always broad, much broader
than a young preglacial valley could have been at that stage of early
youth when its side streams had not cut down to its own depth.
Hence glacial erosion must, under this supposition, be appealed to for
the widening of preglacial canyons, steep-walled and narrow, into the
existing fiord troughs, steep- walled and broad. At the middle of the
fiord troughs, the lateral erosion thus demanded would often measure
thousands of feet, and that in the most massive and resistant crystal-
line rocks.

A second supposition leads to no greater economy of glacial
action. Let it be supposed that the revived streams of preglacial
time had reached maturity before the advent of the glacial period.
In that case, the side streams must have entered the main streams

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DAVIS: GLACIAL EROSION. 291

at accordant grade, and hence the main valleys could not then have
been cut much deeper than the side valleys are now cut ; not so
deep, indeed, for the side valleys have been somewhat deepened by
glacial action, if one may judge by their trough-like form as well as
by the evidence of intense glacial action all over the uplands, even
over most of the surmounting hills and mountains. Hence, to
develop the existing discordant valley system frpm a mature pre-
glacial valley system of normal river erosion, requires a great deep-
ening of the fiords by ice action, again to be measured in thousands
of feet. Thus there seems to be no escape from the conclusion that
glacial erosion has profoundly modified Norwegian topography.
As far as I could judge from my brief excursion over the highlands,
either one of the two suppositions above considered is permissible,
provided only that strong glacial erosion comes after the river work
of the current cycle.

If the Hardanger fiord may be taken as the type of its many
fellows, one may say that hanging lateral valleys are the rule, not
the exception, in Norway. Furtheimore, the smoothed, spurless
walls of the larger fiords, composed of firm bare rock from the
upland to water edge, do not resemble the ravined and buttressed
sides of normal valleys. The marks of downward water erosion
are replaced by what seem to be marks of nearly horizontal pluck-
ing and scouring. Blunt-headed valleys and corries (botner) both
seem beyond production by normal weathering and washing. Yet,
striking as these features are, they do not seem to me so compul-
sory of a belief in strong glacial erosion as the hanging valleys
that have so little relation to the fiords beneath them, and the
flaunting waterfalls that descend so visibly from the hanging
valleys, instead of retiring, as is the habit of falls all over the
un glaciated pai*ts of the world, into ravines where they are hid to
sight from most points of view.

The rocky islands that rise from the shallower parts of the fiords
should not be taken as signs of feeble glacial erosion, but rather as
remnants surviving from the destruction of larger masses in virtue
of some slight excess of resistance. A well-known example of this
kind is near Odde at the head of the large southern arm (Sorfjord)
of the upper Hardanger fiord ; but in the same neighborhood are
several fine hanging valleys, one of which is shown in Plate 3,
Figure Ay its open floor is high above the fiord level ; its cascading
stream, the Strandfoa, descends into Sandven Lake, just south of
the side valley occupied by the well-known Buer glacier.

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292 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

Correlation of River Valleys a7id Olacier Valley a, — Thus far
the consequences of glacial erosion have been described as if thej
were unlike those of river erosion, especially in respect to the pro-
duction of hanging valleys. A just comparison of the two agen-
cies will show that their resemblances are more marked than their
differences, when due allowance is made for their individual
peculiarities.

The likeness of glaciers and rivers has been frequently con-
sidered. The motion of water streams and ice streams is retarded
by bottom and banks, and is fastest in mid-channel where farthest
removed from all hindrances. The motion is faster on strong than
on gentle slopes, and in large than in small streams : the line of
fastest motion departs from the medial axis towards the concave
bank. Forel ('97, 204) and Gannett ('98, 422) have justly com-
pared ordinary valley glaciers, not to rivers that mouth in the sea>
but to rivers that descend from mountains to wither away on pied-
mont deserts. The terminal moraine of the glacier corresponds to
the terminal delta-like fan of withering rivers. The fluctuation of a
withering river following changes of weather or season corresponds
to the secular fluctuations of glaciers, as during the period of about
thirty-five years in the Alps. The advance and retreat at the end
of large glaciers does not occur synchronously with the advance
and retreat of small glaciers, although both large and small glaciers
accomplish their periodic variations of length in the same interval ;
and it is probable that the same contrast obtains in withering^
rivers of different length, although I cannot find any direct state-
ments to this effect. Meunier ('97, 1043) has suggested that certain
peculiaV successions of drift deposits in Switzerland may be the
result of the enlargement of the drift-bringing glacier by the
capture of the head reservoirs of another glacier, after the analogy
of rivers. Gannett and Penck (see abstracts below) have gone
further still and have shown that the hanging valleys, so char-
acteristic of strongly glaciated drainage systems, have a perfect
analogy in the valley systems of ordinary rivers in non-glaciated
areas. This comparison is so instructive that it deserves full state-
ment here.

The ** nice adjustment of declivities " that characterizes the main
and the side valleys of a river system is found only in maturely
developed valleys. The adjustment or accordance between main
trunk and lateral branch obtains only with respect to the surface of
the streams or to the floor of their valleys. The beds of the trunk

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and the branch channels may be discordant at their junction, and
this discordance will increase with the difference in volume of trunk
and branch stream. Truly, the discordance of stream beds is sel-
dom noted, because the beds are hidden by the streams; but if a
river system were laid dry, we may be assured that the beds of the
smaller tributaries would open in the banks of the main river a
number of feet from its bottom. In the case of the Mississippi, the
discordance might easily measure fifty or more feet.

All this applies equally to glacial streams. The surface of a trib-
utary glacier is adjusted to the surface of the trunk glacier that it
joins ; but the depth of the beds may be very different. As long
as the glaciers occupy their channels, the discordance of their beds
may not be often considered, but when a climatic 'change causes
the glaciers to melt away, their channels are called " valleys," and
the discordance of main and lateral glaciated "valley floors" is
taken as an abnormal feature. In reality the discordance is per-
fectly normal to the peculiar system of ice drainage by which it was
produced, however discordant it may be to the system of water
drainage now in possession of the valleys. Let us compare the
maturely developed channels of rivers and glaciers.

ChamieU of Mature Rivers and Glaciers, — A river flows rap-
idly ; and the cross-section of its channel is but a small fraction of
the cross-section of its valley. The river channel is U-shaped, very
broad compared to its depth, while the valley sides flare open,
V-like, above the river banks. The water surface slopes steadily
down-stream, but the channel bed has many small inequalities in
the form of bars and basins, and the water in the bottom of the
basins must ascend a little to get out of them. If the river should
dry away, the deeper parts of the bed would be occupied by pools
of standing water, while the bars would show lines suggestive of
flowing water. The banks of the river channel are smoothly worn
in nearly horizontal lines, parallel to the flow of the river current,
while the sloping sides of a river valley are buttressed with spurs
and scored by the down-hill ravines of descending streams. At the
junction of trunk and branch streams, a moderate discordance in the
level of the channel beds is to be expected ; but this is seldom con-
sidered, because the channels are usually occupied by water and the
beds are hidden.

A glacier moves slowly, and the cross-section of its channel may
be a considerable part of the cross-section of the valley that it
drains. Forel estimates that the glacier of the Rhone, even where

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294 PROCEEDINGS: BOSTON SOCIETY NATURAL fflSTORY.

descending its steep cascade, has only 1 : 12,000,000 of the velocity
of a large river on a similar slop% ('97, 203). The glacial channel is
U-shaped, broad and deep, while the valley flares open, Y-like,
above the ice surface. The ice surface slopes steadily down-stream,
but the bed of its channel is unevenly scoured, here rising in knobs,
there sinking in hollows or basins from which the bottom ice must
ascend a little as it moves forward. When the ice melts away,
lakes occupy the rock basins; the rocky knobs are seen to be
rounded and plucked in a manner suggestive of heavily moving ice.
The banks of the channel are scoured and fluted parallel to the ice
motion ; but above the ice- worn channel the flaring valley sides are
ravined by descending water streams. At the junction of trunk and
branch glaciers a strong discordance in the level of the channel beds
may be expected ; and the discordance becomes conspicuous when
the glaciers melt away and leave their " channels " to be called
"valleys." Hanging side valleys are therefore appropriate as well
as characteristic features of glaciated main valleys. They must
come to be considered even more significant of glacial erosion than
lake basins.

The Cycle of Glacial Denudation, — The points of resemblance
between rivers and glaciers, streams of water and streams of ice,
are so numerous that they may be reasonably extended all through
a cycle of denudation. Let us then inquire if glaciers may not,
during their ideal life history, develop as orderly a succession of
features as that which so well characterizes the normal development
of rivers. The " life history of a glacier *' need not be taken only
in the sense so well illustrated in the last chapter of Russell's " Gla-
ciers of North America," where the glacier is called young when it
is small at the beginning of a glacial climatic epoch; mature when
it is largest during the full establishment of the glacial climate ; and
old when it is vanishing under the re-establishment of a milder cli-
mate. Let us here consider the life history of a glacier under a
constant glacial climate, from the beginning to the end of a cycle of
denudation, just as Russell has considered the "life history of a
river " under a constant pluvial climate, in his " Rivers of North
America." Thus young glaciers will be those which have been just
established in courses that are consequent upon the slopes of a.
newly uplifted land surface ; mature glaciers will be those which
have eroded their valleys to grade and thus dissected the uplifted
surface ; and old glaciers will be those which cloak the whole low-
land to which the upland has been reduced, or which are slowly

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fading in the milder climate of the low levels appropriate to the
close of the cycle of denudation.

Imagine an initial land surface raised to a height of several thou-
sand feet, with a moderate variety of relief due to deformation.
Let the snow line stand at a height of two hundred feet. As ele-
vation progresses, snow accumulates on all the upland and highland
surfaces. Glaciers are developed in every basin and trough ; they
creep slowly forward to lower ground, where they enter a milder
climate (or the sea) and gradually melt away. At some point
between its upper heads and its lower end, each glacier will have a
maximum volume. Down stream from this point, the glacier will
diminish in size, partly by evaporation but more by melting ; and
the ice water thus provided will flow away from the end of the
glacier in the form of an ordinary stream, carving its valley in
normal fashion. Some erosion may be accomplished under the
upper fields of snow and n^v^, but it is believed that more destruc-
tive work is done beneath the ice. The erosion is accomplished by
weathering, scouring, plucking and corrading. Weathering occurs
where variations of external temperature penetrate to the bed-rock,
as is particularly the case between the s^racs of glacial cascades,
and again along the line of deep crevasses or bergschrunds that are
usually formed around the base of reservoir walls, which are thus
transformed into corries (cirques, karen, botner) as has been sug-
gested by several observers; scouring is the work of rock waste
dragged along beneath the glacier, by which the bed-rock is ground
down, striated and smoothed ; plucking results from friction under
long-lasting heavy pressure, by which blocks of rock are removed
bodily from the glacier bed and banks ; corrading is the work of
subglacial streams, which must be well charged with tools, large and
small, and which must often flow under heavy pressure and with
great energy. All these processes are here taken together as " gla-
cial erosion."

Let it be assumed that at first the slope of a glacier's path was
steep enough to cause it to erode for the greater part or for the
whole of its length. Each young glacier will then proceed to cut
down its consequent valley^ at a rate dependent on various fac-
tors, such as depth and velocity of ice stream, character of rock
bed, quantity of ice-dragged waste, and so on ; and the eroded

^A valley is understood to Include the channel that is eroded along its floor. The
channel, with its bed and banks, Is therefore that i)art of a valley which is occupied by
the stream.

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296 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORT.

channel in the bottom of the valley will in time be given a depth
and width that will better suit the needs of ice discharge than did
the initial basin or trough of the uplifted surface. The upper slopes
of the glacial stream will thus be steepened, while its lower course
will be given a gentler descent. Owing to the diminution of the
glacier toward its lower end, the channel occupied by it will dimin-
ish in depth and breadth downwards from the point of maximum
volume; this being analogous to the decrease in the size of the
channel of a withering river below the point of its maximum volume.
A time will come when all the energy of the glacier on its gentler
slope will be fully taxed in moving forward the waste that has been
brought down from the steeper slopes ; then the glacier becomes
only a transporting agent, not an eroding agent, in its lower course.
This condition will be first reached near the lower end, and slowly
propagated headwards. Every part of the glacier in which the
balance between ability to do work and work to be done is thus
struck may be said to be " graded " ; and in aU such parts, the sur-
face of the glacier will have a smoothly descending slope. Maturity
will be reached when, as in the analogous case of a river, the nice
adjustment between ability and work is extended to all parts of a
glacial system. In the process of developing this adjustment, a
large trunk glacier might entrench the main valley more rapidly
than one of the smaller branches could entrench its side valley ;
then for a time the branch would join the trunk in an ice-rapid of
many s^racs. But when the trunk glacier had deepened its valley
so far that further deepening became slow, the branch glacier would
have opportunity to erode its side valley to an appropriate depth,
and thus to develop an accordant junction of trunk and branch ice
surfaces, although the cJuinnds of the larger and the smaller streams
might still be of very unequal depth, and the channel beds might
stand at discordant levels. If the glaciers should disappear at this
stage of the cycle, their channels would be called valleys, and the
discordance of the channel beds might naturally excite surprise.
The few observers who, previous to 1898, commented upon a dis-
cordance of this kind, explained it as a result of excessive erosion of
the main valley by the trunk glacier; while the hanging lateral
valleys were implicitly, if not explicitly, regarded as hardly changed
from their preglacial form.

When the trunk and branch glaciers have developed well-defined,
maturely graded valleys, the continuous snow mantle that covered
the initial uplands of early youth is exchanged for a discontinuous

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cover, rent on the steep valley sides where weathering comes to
have a greatly increased value, and thickened where the ice streams
have established their courses. This change corresponds to that
belween the ill-defined initial drainage in the eai-ly youth, and the
well-defined drainage in the maturity, of the river cycle.

It is probable that variations in rock structure will have permitted
a more rapid development of the graded condition in one part of
the glacial valley than in another, as is the case with rivers of water.
Steady-flowing reaches and broken rapids will thus be produced in
the ice stream during its youth ; and the glacial channel may then
be described as " broken-bedded." But all the rapids must be worn
down and all the reaches must become confluent in maturity. It is
eminently possible that the reaches on the weaker or more jointed
rocks may be eroded during youth to a somewhat greater depth
than the sill of more resistant or less jointed rock next down stream ;
and if the glacier should vanish by climatic change while in this
condition, a lake would occupy the deepened reach, while the lake
outlet would flow foi-ward over rocky ledges to the next lower reach
or lake. Many Norwegian valleys today seem to be in this con-
dition. Indeed some observers have described broken-bedded val-
leys as the normal product of glacial erosion, without reference to the
early stage in the glacial cycle of which broken-bedded glacial chan-
nels seem to be characteristic. Truly, it is not always explicitly
stated that the resistance of the rock bed varies appropriately to the
change of form in a broken-bedded channel ; but the variations of
structural resistance or firmness that the searching pressure and fric-
tion of a heavy glacier could detect might be hardly recognizable to
our superficial observations ; and on the other hand the analogy of
young ungraded glaciers with young ungraded rivers seems so nat-
ural and reasonable that broken-bedded glacial channels ought to be
regarded only as features of young glacial action, not as persistent
features always to be associated with glacial erosion. If the glaciers
had endured longer in channels of this kind, the " rapids " and
other inequalities by which the bed may be interrupted must have
been worn back and lowered, and in time destroyed.

If a young glacier erodes it« valley across rocks of distinctly dif-
ferent resistances, a strong inequality of channel bed may be devel-
oped. Basins of a considerable depth may be excavated in the
weaker strata, while the harder rocks are less eroded and cross the
valleys in rugged sills. Forms of this kind are known in Alpine
valleys ; for example, in the valley of the Aar above Meiringen

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(Wallace, '96, 176) and in the lower Gastemthal near ite junctioii
with the Kanderthal ; in both these cases the basins have been
aggraded and the sills have been trenched by the postglacial streams.
In the lower Gastemthal the height and steepness of the rocky
sill, when ai)proached from upnstream, is astonishing ; its contrast to
the basin that it encloses is difficult enough to explain even for those
who are willing to accept strong glacial erosion. It should, how-
ever, be noted that river channels also are deeper in the weaker
rocks up-stream from a hard rock sill ; if the river volume should
greatly decrease, a small lake would remain above the sill, drained
by a slender stream cutting a gorge through the sill.

If an initial depression occurred on the path of the glacier, so
deep that the motion of the ice through it was much retarded, an
ice-lake would gather in it. Then the waste dragged into the basin
from up-stream might accumulate upon its floor until the depth of
the basin was sufficiently decreased and the velocity of the ice
through it sufficiently increased to bring about a balance betw-een
ability to do work and work to be done. Here the maturely graded
condition of the ice stream would have been attained by aggrading
its bed, instead of degrading it ; this being again closely analogous
to the case of a river, which aggrades initial depressions and degrades
initial elevations in producing its maturely graded course.

Water streams subdivide toward the headwaters into a great
number of very fine rills, each of which may retrogressively cut its
own ravine in a steep surface, not cloaked by waste. But the
branches of a glacial drainage system are much more clumsy, and
the channels that they cut back into the upland or mountain mass
are round-headed or amphitheatre-like ; but the beds of the branch-
ing glaciers cannot be cut as deep as the bed of the large glacial
channel into which they flow : thus corries, perched on the side-
walls of large valleys, may* be produced in increasing number and
strength as glacial maturity approaches, and in decreasing strength
and number as maturity passes into old age. As maturity ap-
proaches, the glacial system will include not only those branches
that are consequent upon the initial form, but certain others which
have come into existence by the headward erosion of their n6v6
reservoirs following the guidance of weak structures ; thus a ma-
turely developed glacial drainage system may have its subsequent
as well as its consequent branches. It is entirely .conceivable, as has
been suggested by Meunier, that one ice stream may capture the
upper part of another. The conditions most favorable for such a

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process resemble those under which river diversions and adjustments
take place; namely, a considerable initial altitude of the region,
allowing a deep dissection ; a significant difference of drainage areas
or of slopes, whereby certain glaciers incise deeper valleys than
others ; a considerable diversity of mountain structure, permitting
such growth and arrangement of subsequent glaciers as shall bring
the head reservoir of a subsequent ice stream alongside of and
somewhat beneath the banks of a consequent ice stream. Thus
glacial systems may come to adjust their streams to the structures
upon which they work, just as happens in river systems.

The load transported by a glacial system may at first be supplied
largely by waste plucked and scoured from the beds of the glacial
channels as well as by waste detached from the enclosing slopes ;
but in time, when the graded condition of the chief channels is
reached and their further deepening almost ceases, by far the largest
share of load will be supplied from the subaerial valley sides, where
weathering of the ordinary kind will ravine the slopes, thus produc-
ing a topography that is strongly contrasted with the smooth walls
of the glacial channels. If the initial glacial system should incise its
channels so deeply beneath a lofty highland that the supply of waste
from the valley sides continued to increase after the development of
graded glacial channels, it is conceivable that the channel- beds
might have to be aggraded for a time, as is believed to be the case
\^4th river channels under similar conditions; but owing to the
receipt on the glacial surface of waste from the valley sides, it is also
conceivable that this analogy may not closely obtain. Toward the
end of the ice stream it may well happen that the diminution of its
volume and the consequent diminution of its capacity to do work
will result in the aggradation of its bed by waste that cannot be
carried further forward. At the same time, the outflowing river may
be unable to wash away all the waste that is delivered to it, and so, for
a time through later youth and early maturity, the river may act as
an aggrading agent and build up a broad, flat alluvial fan, such as
fronts the terminal moraines of the Alpine glaciers that once
descended to the plain of Lombardy. Some response to the change
thus produced in the altitude of the end of the glacier may be
expected far up its channel, whose bed would thus come to be
aggraded with till. Similarly, the ice sheets that spread from the
Scandinavian and Laurentian highlands over the lowlands on the
south changed their behavior from degrading agents in the central
area to aggrading agents on the peripheral area. Hence, a belief in

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300 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

effective erosion is not antagonistic to a belief in effective deposition
in the case of glaciers any more than in the case of rivers. In each
case the action varies appropriately to its place in the drainage
system and to its stage in the cycle. But there will be a later stage,
when the wasting of the superglacial slopes reduces them to mod-
erate declivity, so that the waste delivered from them decreases in
quantity ; then the outflowing water stream at the end of the glacier
may become a degrading agent ; the altitude of the end of the gla-
cier may be slowly lessened ; and a very slow and long-continued
deepening of the whole glacial channel will take place, without
requiring a departure from an essentially graded condition.

As the general denudation of the region progresses, the snow fall
must be decreased and the glacial system must shrink somewhat,
leaving a greater area of lowland surface to ordinary river drainage.
When the upland surface is so far destroyed that even the hill tops
stand below the 200-foot contour, the snow fields will be represented
only by the winter snow sheet, and the glaciers will have disap-
peared, leaving normal agencies to complete the work of denudation
that they have so well begun.

If a snow line at sea-level be assumed, glaciation would persist
even after the land had been worn to a submarine plain of denuda-
tion at an undetermined depth beneath sea-level. The South Polar
regions offer a suitable field for the occurrence of such a surface.

Whether glaciers of the Norwegian or of the Alpine type shall
occur, is dependent partly on initial conditions, partly on the stage
of advance through the cycle of denudation. If the initial form
offer broad uplands, separated by deep valleys, snow fields of the
Norwegian type may have possession of the uplands during the
youth of the glacial cycle ; but when maturity is reached, the up-
lands will be dissected, and the original confluent snow field will be
resolved into a number of head reservoirs, separated by ridges. On
the other hand, as the later stages of the cycle are approached, the
barriers between adjacent reservoirs will be worn away, and they
will tend to become confluent, here and there broken only by
Nunatuker. If the snow line lay low enough, a completely con-
fluent ice and 8no\<^ shield would cover the lowland of glacial denu-
dation when old age had been reached. If the glacial conditions of
Greenland preceded as long as they have followed the glacial period
over the rest of the North Atlantic region, who can say how far the
ice of the Greenland shield has modified the forms on which its
work began I

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Glacial Distributaries, — If a maturely dissected mountain range
were occupied by snow-fields and glaciers of large size, certain
peculiar results might be expected near the mountain base. Under
normal preglacial conditions, a small low ridge sufiices for the com-
plete separation of two river systems, because the channels of rivers
are so small in comparison to their valleys. But glacial channels are
a large part of their valleys, and when great glaciers from the lofty
mountain centres descend by the master valleys to the mountain
flanks or even to the piedmont plains, distributary ice streams or
outflowing branches may naturally enough be given off wherever
the ice surface rises high enough to overtop the ridges by which the
master valleys are separated from adjacent minor valleys. If a dis-
tributary branch has suflicient strength and endurance, it may wear
down the ridge that it crosses and thus increase and perpetuate its
lateral discharge ; but it cannot usually be expected to erode a
channel as deep as that of the main glacier from which it departs.
On the disappearance of the ice, a hanging valley will be left above
the floor of the master valley ; but in this case, the drainage of the
hanging valley will be away from, not toward, the master. Here
we probably have the explanation of those broad hanging valleys
which lead from the valley of Lake Maggiore on the west and, less
distinctly, from that of Lake Como on the east to the compound
basin of the intermediate Lake Lugano. On going southward by
rail from BeUinzona to Lugano, along a stretch of the St. Gotthard
route between the great tunnel and Milan, the railway obliquely
ascends the southeastern wall of the trough-like valley of the Tieino
just above the head of Lake Maggiore ; and at a height of several
hundred feet over the delta flood-plain the line turns off to a well-
marked hanging valley in which the stream runs away from the
Tieino to Lake Lugano. The notch made by this supposed glacial
distributary is a conspicuous feature in the view from BeUinzona
and thereabouts.

The anomalous forking of Lake Como and the open branch from
the main valley of the Rhine at Sargans through the trough of
Wallen See to Lake Zurich appear to be the paths of large glacial
distributaries which eroded their channels deeply across divides that
presumably existed in preglacial time. The west wall of the main
valley of the Is^re in the Alps of Dauphiny, southeastern France, is
deeply breached by passes that lead northwest to the troughs of
Lakes Annecy and Bourget, through which the distributaries of the
Is^re glacial system must have flowed. Lug^on ('97, 62-70) has

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302 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

explained the breaches as marking the former northwest paths of
transverse members of the Is^re system, from which they have been
diverted by the subsequent growth of the main longitudinal valley —
that northeast- southwest part known as the Gr^sivaudan — above
Grenoble. HLs discussion of the problem takes, however, no account
of modifications of valley depth by glacial erosion ; and as this must
have been considerable (for the valleys hereabouts have superb
basal cliffs, as appears in the valley of the Romanche by Bourg
d'Oisans in Plate 3, Fig. J5, after a photograph by Neurdin Fr^re^
of Paris), it may well be that the rearrangement of river courses
in this interesting region is not altogether the work of river action.
Similarly, the various modifications of the Rhine system in eastern
Switzerland, explained by Ileim as the work of streams alone, may
come to be at least in part referred to ice erosion.

It may be further supposed that if the preglacial valleys were
so arranged that a glacial distributary found a shorter and steeper
course to the piedmont plain or to the sea than that followed by the
master glacier, the distributary^ might under a long enduring glaci-
ation become the main line of glacial discharge ; and if so, it could
be eroded to a greater depth than the former master valley at the
point of divergence. In such a case, the postglacial river drainage
would differ significantly from the preglacial. There is reason for
believing that examples of this kind are to be found in Norway, the
evidence of which will soon be published in an essay by Barrett
(1900). The diversion of the head of a stream in the Sierra Costa
of northwestern California to a deei)er-l\ing valley through a
gorge cut by a glacial distributarj^ has lately been described by
Hershey (1900, 47).

The Depth of Mature Glacial Chaiinels, — The depth with
respect to sea-level to which the channels of a glacial system may be
eroded when the graded condition is reached, is a subject of special
interest. ' For many miles along the lower course of a branchless
trunk glacier, its volume is lessened by melting and evaporation,
and at its end the ice volume is reduced to zero ; slow ice motion
being progressively replaced by rapid water motion. In such a case
the law of continuity does not demand that the ice velocity shall be
inversely proportional to the area of the cross section, as is the case
in the normal river (where it is assumed that there is no loss by
evai)oration). Indeed, in the lower trunk of a mature glacier, it
may well be that the velocity of ice movement is in a rough way
directly proj)ortional to cross-section area. This appears to be veri-

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DAVIS: GLACIAL EROSION. 303

fied by meafluremente of the Rhone glacier, where the mean annual
movement is 110 met. in the heavy trunk above the cascade, 27
met. just below the cascade, and only 5 met. close to the melting
front (Forel, '97, 203) . Evidently then, the erosion of the glacial
bed, in so far as it is determined by the pressure and motion of the
ice stream, will have its maximum some distance up-stream from the
end of the glacier (J. Geikie, '98, 236). The glacial channel must
therefore become narrower and shallower as its end is neared, as has
already been stated. If the glacier ends some distance inland from
the sea, its action will be conditioned by the grade and length
of the river that carries away the water and waste from its lower
end. The deepening of the distal part of the channel accomplished
in youth might be followed by a shallowing for a time during matu-
rity, when the accumulation of mor^nal and washed materials in
front of the glacier compelled its end to rise. Now it may well be
conceived that the surface slope of such a glacier near its end is less
than the angle between the surface and the bottom of the glacier ;
and in this case, the glacial floor must become lower and lower for
a certain distance up-stream. If such a glacier should. melt away,
the distal part of its channel would be occupied by a lake, although
even the head of the lake may not reach to the locus of maximum
glacial erosion. Lakes Maggiore, Como and Garda seem to occupy
basins whose distal enclosure by heavy moraines and sheets of over-
washed gravels has added to the depth produced by erosion further
up-stream. It would ^eem, however, that a lake basin thus situated
must be only a subordinate incident in the general erosion of the
whole length of the glacial channel. Too much attention has, as a
rule, been given to lakes of this kind, and not enough to the other
effects of glacial action ; it seems especially out of proportion to sup-
pose that the maximum erosion by a glacier takes place near its end,
as has been done by some authors, on account of the prevalent
occurrence of lakes in this situation.

If a glacier advances into the sea and ends in an ice cliff, from
which ice blocks break off and float away, something of a basin-like
form of its lower channel may be produced ; but the dimensions of
this basin will be determined by the climate at the termination of
of the glacier. If the climate is such as to allow the glacier to enter
the sea in maximum volume, then a basin is not to be expected. The
more the glacier diminishes towards its end, the less erosion and the
more deposition may occur beneath it, and the more of a basin may
be developed inland from its end.

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The depth to which a glacier may cut its channel when it enters
the sea is of particular importance. If the glacier is a thousand feet
thick at its end, it must continue to press upon and scour its bed
until only about 140 feet of ice remain above sea4evel ; ita channel
will thus be worn more than 800 feet beneath sea-level. Truly, the
latter part of this work will be performed with increasing slowness ;
but if time enough be allowed the work must be accomplished, just
as is the case with rivers. If a glacier should melt away from its
deep entrenchment, its channel would be occupied by an arm of the
-sea or fiord, reaching many miles into the land. The fiord might be
shallower at its mouth than further inland, if differential erosion and
deposition had occurred along its channel. Yet even this result is
analogous to the case of a river ; for if the Mississippi were to dis-
appear in a prolonged drought, a slender arm of the sea would invade
the river channel many miles up-stream from the delta front. Indeed,
the Mississippi offers an excellent example of a channel that is ba-
sined inward from the river mouth ; for while it is only a score of
feet deep at the passes where most of its sediment is deposited, it is
several score of feet in depth further up-stream ; and the slender
arm of the sea that would occupy its channel if it should disappear
by climatic change, would be truly fiord-like in having a less depth
at its mouth than further inland.

An important corollary from this conclusion — perhaps not so much
of a novelty to glacial erosionists as to their confreres of the opposite
oi)inion — is that the depth of water in the fiords of a strongly gla-
ciated coast is not a safe guide to the movement of the land since
preglacial time. If there had been a still-stand of the earth's crust
through the whole glacial period, the preglacial river channels that
were graded down a little below sea-level at their mouths would be
replaced by glacial channels that might be eroded hundreds of feet
below sea-level. The depth of fiords thus seems to depend on the
size of their ancient glaciers, on the height of the mountain back-
ground, and on the duration of the glacial period, as well as on
movements of the land. If liberal measures of glacial erosion and
glacial time are allowed, no depression of glaciated coasts sinc^ pre-
glacial time' is needed to account for their peculiar features. The
glacial channels may have been simply invaded by the sea, as the
ice melted away, without any true submergence.

Even the advocates of strong glacial erosion do not seem to have
expUcitly recognized the full importance of this possibility. James
Geikie, for example, writes : " The fiords of high latitudes and the

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DAVIS: GLACIAL EROSION. 305

narrow inlets of non-glaciated lands are simply submerged land-
valleys; the intricate coast-lines of such regions have been deter-
mined by preceding subaerial denudation." Again: "In a word,
fiords are merely the drowned valleys of severely glaciated moun-
tain-tracts." ('98, 263, 250.) The deep waters in the valley of the
Hudson through the Highlands of southeastern New York are the
most fiord-like in the eastern United States, and they are universally
explained as the result of submergence of a normal river valley ; but
the constricted ice current that must have flowed through the High-
land gorge may have been energetic enough to deepen its bed
beneath sea-level, and since the ice melted away, who can say how
much submergence beneath preglacial levels has taken place. I do
not know how far this view of the matter has been taken by ear-
lier advocates of strong glacial erosion, but for my own part, the
acceptance of such a possibility means a complete reversal of the
belief that I held two years ago. The reversal is, however, accom-
panied by the memory that it was always difficult to understand
why submergence and glaciation were so closely associated : even if
glaciation had caused depression, it was diflicult to understand why
the relief from ice pressure in postglacial time had not now been
followed by a rise of the land much nearer to its former altitude
than would be the case if the greater part of the depth of fiords is
explained by submergence.

The Origin of Carrie Hasins. — On pursuing the above line of
consideration a little further, it may give some light on the occur-
rence of the small rock basins that are so often found in the floor of
cliff-walled corries. Imagine that a large glacial system has become
maturely established, and that it " rises " in many blunt head-
branches that have excavated corries in a preglacial mountain mass,
and have cut down channels, at their junction with the larger
branches or trunk glacier, to a depth appropriate to their volume.
Unless the erosion of the corries has been guided by differences of
rock structure, there does not seem to be reason for their possessing
a basined floor at this stage of development; but if a change of
climate should now cause the trunk glacier to disappear, while many
of the blunt head-branches remain in their corries, each little glacier
thus isolated will repeat the conditions of erosion above inferred for
the tnink glacier ; and if this style of glaciation linger long enough,
rock basins may very generally characterize the floors of the corries
when the ice finally melts away. Figure 7 may make this clearer.
Let the broken line, ABC, be the slope of a preglacial lateral ravine

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306 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

which reaches a trunk stream at C, while ADC is the profile of an
adjoining lateral spur. After vigorous and mature glaciation, the
dotted line, GE, may represent the surface slope of a lateral glacier,
and GHJ that of the lateral gla<*ier bed ; while EFL is the surface
of the trunk glacier, and EKL the bed. The lower part of the
lateral spur has been cut off to make the basal cliff beneath D. On

Fig. 7. Diagram-section of a lateral valley with a corrie basin.

the disappearance of the trunk glacier at this stage, the shrunken
side glacier, GNJH, occupies its corrie or hanging valley, which
opens at J on the oversteepened wall, DJK, of tlie evacuated channel
of the trunk glacier. Let the maximum erosion of the corrie glacier,
as conditioned by pressure and motion, be at H. Then after some
time the weathering of the cliff walls and the erosion of the floor
will have transformed the corrie and its glaoier to a form,
G'N' J'H', such that the deepening of the glacial bed should be a
maximum at HIP. The continuous slope of the glacial bed, GHJ,
appropriate to the time when the lateral glacier joined the trunk
glacier, may thus be transformed into a basined curve, G'H'J',
appropriate to a small glacier terminating at J' ; and on the disap-
pearance of the small glacier, a tarn or rock-basin lake may occupy
the depression at II'. It is on the basis of a supposition like this
that a determination has been attempted of the altitude at which
the shrinking remnants of an extensive glacial system endured for
a time before their entire disappearance (J. Geikie, *98, 233).
Richter's supposition that the uplands of Norway result from the

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DAVIS: GLACIAL EROSION. 307

consumption of pre-existent mountains by the great extension of
corrie-glacier floors, each similar to J' H', thus seems mechanically
possible ; but it is nevertheless climatically very improbable, and it
seems to me deficient in not attributing enough work to normal pre-
glacial erosion.

Overdeepened Valleys and Oversteepened Walls, — As in the
case of the normal cycle of denudation in which the life history of
river systems is involved, so in the glacial cycle, all manner of com-
plications may arise, causing great departures from the ideal case.
The assumed initial land form may be a surface previously more or
less dissected by river erosion, on which glaciers must then proceed
to develop a drainage system appropriate to their own peculiar
needs, as has been partly considered above in connection with glacial
distributaries. It will be instructive to make out a good series of
examples illustrating different combinations of river and glacial
action, and including young, mature and old river valleys, modified
by young or mature glaciation. For example, the existing vaUey
of the Rhue in the Central Plateau of France shows a submature
river valley with incised meanders, moderately affected by young
and relatively light glaciation ; the valley of the Ticino in the south-
ern Alps is a well-matured preglacial river-valley system, modified
by strong submature glaciation. The fiords of Norway result from
the submature and intense glaciation of a river-valley system whose
stage of preglacial development is not yet well determined.

Interruptions of regular progress in the glacial cycle must, as in
the river cycle, be occasioned by elevation, depression, or deforma-
tion of the land mass ; but no examples of complications of this kind
can be adduced. Variations of climate may replace creeping glaciers
in young, mature or old stages of development, by flowing rivers ;
and the early stages of such rivers are of much importance among
existing geographic forms. Lakes, delaying the river flow, occupy
the depressions of the glaciated surface, as has been known since
Ramsay first pointed out the correlation of lacustrine and glaciated
regions in 1861 ; but the analogy between lakes in the beds of
melted glaciers and pools in the beds of dried-up rivers has perhaps
not been suiRciently insisted upon. Waterfalls connect the streams
that occupy the discordant beds of glacial channels, as has lately
been clearly set forth. Landslides frequently occur after the sup-
porting glacier withdraws from the oversteepened banks of its huge
channel ; fallen masses of this origin have been repeatedly mistaken
for moraines in Alpine valleys, as has been lately shown by Brtlck-

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308 PROCEEDINGS: BOSTON SOCIETY NATURAL mSTORY.

ner. Every lake or fiord is an effective lowering of base-level for
the stream above it ; for the level of a body of standing water is
essentially the same at both ends. As fast as the inflowing river
builds its delta forward at the head of the lake or fiord, its flood
plain must rise up-stream and aggrade the valley floor. This process
is very j)ronounced in many Alpine valleys, where the aggraded
valley floor has a relatively rapid descent on account of the plentiful
and coarse detritus furnished by the active side streams. Indeed,
every ravine furnishes a great quantity of rock waste, whose descent
is analogous to rej)eated landdides of small dimensions. The valley
floor beneath the ravines is invaded by great alluvial fans, and the
main stream is driven away toward the further valley wall by their
rapid advance. At every floo<l, the waste 8uj)j)lied from the fans is
swept abundantly into the main stream, whose flood-plain grows
rapidly as a delta in the upper end of each lake tliat it enters. The
delta of the Ticino seems to have advanced so far into what was
originally the basin of Lake Maggiore that the apparent height of
the hanging lateral valleys steadily decreases toward the lake ; and
for several miles above the head of the lake the lateral valleys seem
to enter the main valley almost at grade, although there can be little
doubt that if all the delta alluvium were removed, the lateral valleys
would be found to stand high above the rock floor of the main val-
ley. The standing lakes, the aggrading flood-plains, and the grow-
ing fans all show that the bed of the glacial channel has been worn
too deep to serve as a valley floor for the existing river ; the river
must aggrade, with water or with waste, the bed of the channel that
the glacier degraded; hence Penck has suggested that glaciated
valleys of the Alpine kind should be called "overdeepened." In
the same way, the waterfalls from the hanging valle}"^, the shower-
ing waste that forms the fans, and the landslides from the basal
cliffs, all show that the banks of the glacial channel — the lower
walls of the existing valleys — are too steep ; and they may be
therefore called " ovei-steepened." Much glacial work had to be
done uj)on the mature preglacial valleys of river erosion, to bring
them into mature adjustment with the needs of glaciers ; much river
work must likewise now be done upon the overdeepened glaciated
valleys, and upon their oversteepened walls and their hanging
branches, before they can be maturely adjusted again to the needs
of rivers.

Practical Utility of the Ideal Glacial Cycle. — In every case, the
full understanding of the conditions developed by any system of

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DAVIS: GLACIAL EROSION. 309

glaciers, existing or extinct, can be reached only by a complete
analysis of the conditions under which they began to work, of the
energy with which they worked, of the part of a cycle during which
they worked, and of the complications of climatic change or of
crustal movements by which their work was modified in this way or
in that. A partial analysis may suffice for a particular instance ; but
the explorer will be better equipped for the explanation of all the
instances that he discovers if he sets out with a well-elaborated con-
ception of the ideal glacial cycle of denudation, and of the compli-
cations it is likely to suffer. However extensive and definite this
conception may be, exploration will probably require its further
extension and definition ; however brief exploration may be, it will
probably be aided by an orderly examination of all pertinent knowl-
edge previously accumulated.

As a practical instance of the value of the glacial cycle, we may
consider the aid given toward the solution of certain problems by
the careful reconstruction — or at least the conscious attempt at
reconstruction — of the form of the land surface on which the pleis-
tocene glaciers began their work, and by the legitimate deduction of
the characteristics of maturity in the cycle of glacial erosion. Beyond
the mature stage, we may seldom have occasion to go, as there do
not seem to be actual examples of more advanced glacial work. The
initial form on which pleistocene glacial action began is in no case
known to be that implied in the opening paragraphs of the section
on the Glacial Cycle ; namely, a land mass freslily uplifted from
beneath the sea and not previously carved by the streams of an
ordinary or normal cycle of erosion. In central France, for example,
the initial form was an uplifted and submaturely dissected peneplain,
in which valleys with incised meanders had been habitually devel-
oped. It was there of the greatest assistance to carry into the
glaciated area a clear picture of its preglacial form, as determined by
generalizing the adjacent non-glaciated area. At the same time, the
ideal picture of a maturely developed glacial drainage system, with
smooth-sided troughs, was seen to represent a much more advanced
condition than was attained in the rugged valley of the Rhue ; and
thus a tolerably definite idea was gained of the youthful stage of
glacial development, somewhere between its beginning and its
maturity, and of the amount of destructive work needed to reach
this youthful stage. This elementary example illustrates a method
embodying the cycle of glacial denudation that ought to be applied
whenever possible.

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The larger Nom egian fiords may be instanced as glacial channels
that present every appearance of having advanced far toward the
mature stage of a cycle of glacial denudation from an initial or pre-
glacial form not yet well understood. The variation of form
between the main fiords and their branches gives some indication
that the glacial work waa accomplished in several successive epochs,
with the interglacial epochs of normal river work between ; but this
is only a suggestion, needing much more field work before it can be
assured. Not only the deep fiords, but the hanging valleys and the
uplands also, have been ice-scoured ; for hanging valleys frequently
have a well-defined U-section, and sometimes receive secondary
hanging valleys from the enclosing uplands; and the streams of
the uplands exhibit rej)eated departures from the forms of normal
erosion. Although possessing little drift, the uplands frequently
bear lakes of moderate depth and irregular outline ; in 8]f)ite of the
breadth to which the upland valleys are opened between tiie sur-
mounting hills and mountains, their streams frequently change from
wandering at leisure in split or braided channels along broad floors,
to dashing down in haste over rocky rapids : a behavior that is mani-
festly inconsistent with that of the mature drainage of a normally
denuded region. Even the surmounting hills exhibit strong scour-
ing on their up-ice-stream side. It does not therefore seem permis-
sible to conclude that the hanging valleys which open on the walls of
the greater fiords have not been deepened by ice erosion because
they escaped the more severe glaciation that scoured out the fiords
themselves. All the valleys have been glaciated, and all have been
significantly modified from their preglacial form. The discordance
of overdeepened main fiord and hanging lateral valley seems to me
best explained as the result of the mature development of glacial
drainage, in which the chief trunks and the larger branches of the
glacial systems had for the most part reached a graded condition.
Trunk and branch glaciers would then have united at even grade
as to their upper surface, and the trunk and branch channels would
have had dimensions satisfactory to the ice currents which flowed
through them, but the channel beds would have been discordant, as
they are found to be.

Review of Previous Writings oar Hanging Valleys.

It has already been stated that hanging side valleys and over-
deepened main valleys have not yet been generally given the impor-

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DAVIS: GLACIAL EROSION. 311

tance that they deserve as witnesses to strong glacial erosion.
Russell, in his "Glaciers of North America" (1897), makes no
mention of discordant lateral and main valleys when discussing
glacial erosion. James Geikie, in his "Earth Sculpture" (1898),
allows to discordance of glaciated valleys hardly more than a second-
ary importance in abstracts and quotations from Wallace's accounts
of Alpine lakes and from Richter's essay on Noi-way (see below),
while the glacial erosion of lake basins is much more fully treated.
Yet of all the facts that point to strong glacial erosion, none seem
to give testimony so unanswerable as do hanging valleys. The fol-
lowing extracts will serve to illustrate the gradually increasing
recognition of their importance.

Forbes on the Waterfalls of Norway. — Thinking that some
interesting early observations on the hanging valleys of Norway
might be recorded in Forbes' book of travels in that countiy, I
looked up waterfalls in his index and there found a reference to the
cause of their profusion, which was stated as follows.

"The source of this astonishing profusion of waters is to be
found in the peculiar disposition of the surface of the country so
often referred to. The mountains are wide and flat, the valleys are

deep and far apart As the valleys ramify little .... and are

wholly disconnected from the fields [uplands] by precipitous slopes,
it follows that the single rivers which water ' those valleys .... are
supplied principally by streamlets which, ha^dng run long courses
over the fields^ are at last precipitated into the ravines in the form
of cascades." ('53, 251).

Forbes was an excellent observer, yet this quotation is about
equivalent to saying that there are many waterfalls in Norway
because there are steep slopes over which the streams of the uplands
must descend. The quotation deserves a place here if for nothing
more than to show the advance of a half century in regard to what
constitutes the cause of a geographical feature.

Mc Gee on ^filacial Canyons^ 188S. — The earliest article that I
have found touching on this subject is the brief abstract of a paper
read by McGee before the American Association in 1883, entitled
" Glacial Canons." Observations in the Sierra Nevada led this keen
observer to state that ** the effect of the temporary occupancy of a
t}7)ical water-cut canon by glacier ice will be to (1) increase its
width, (2) change the original V to a U cross profile, (3) cut off the
terminal portions of tributaiy canons and thus relatively elevate
their embouchures, (4) intensify certain irregularities of gradient in

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the canon-bottom, (5) excavate rock-basins, (6) develop cirques,
and, in general, transform such canon into an equally typical glacial
canon '* ('83, 238). A later paper by the same author is referred
to below.

Hanging VaUeya in the Alps, — Valleys that are here called
*' hanging " have frequently been described by observers in the Alps,
but either without particular reference to their discordant relation to
the main valleys, or with acceptance of normal erosion in the main
valley as the cause of their discordance. Rtttimeyer's and Heim's
views on discordant lateral valleys have been already referred to.

An account of the Salzachthal in the Tyrol, by Brttckner, describes
it as one valley in the bottom of another; the deeper one being
relatively narrow and steej)-sided, wliile the sides of the higher
valley flare wide open ; the side streams are described as falling into
the deeper main valley ; but tliis significant feature is not mentioned
as if it were of general occurrence, nor is it explained by glacial
erosion ('85, 95).

Lubbock, in his ** Scenery of Switzerland," follows Rtitimeyer
and Heira. After stating that the side valleys of the Reuss have a
moderate grade which brings them out at the level of one of the ter-
races or Thalstufen of the main valley, from which their streams cas-
cade down into the main stream, Lubbock writes : " It is obvious that
this terrace represents a former * Thalweg ' of the Reuss with much
less fall- than it has now, and that the river has deepened its valley
more rapidly than the lateral streams, so that these glens open at
some (listanee up the side of the valley, and their waters join the

Reuss by rapids or waterfalls The valley shows clear evidence

of glacial action. The hard rocks are in places quite polished. This
is especially the case with the buttresses which stand like doorposts
where the lateral glens open into the main valley, and particularly
on the right side of the eastern glens, tlie left of the western, where
of course the pressure of the ice was greatest '* ('96, 332, 334) .

Russell on Hanging Valleys^ 1887, — Russell gave a detailed
account of what may be called hanging valleys in his report on the
Quaternary Histoiy of Mono Valley, California. In a section on
"high lateral canons," he says: "In a number of instances in the
Mono basin the low-grade glaciated canons receive branching caiions
at a considerable elevation above their bottoms, the branches also
having a low grade. This is illustrated where Lake Canon opens
into Lundy Canon. Each of these gorges has an approximately
horizontal bottom near the place of union, but the former is a thou-

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DAVIS: GLACIAL EROSION. 313

sand feet higher than the latter. The stream flowing through Lake
Canon descends precipitously over a rocky face in order to join
Lundy Creek. The bottom of the higher canon is about on a level
with the main lateral moraine in the lower caiion. The same series
of phenomena is repeated where Silver Creek descends over a rocky

face to join Rush Creek It might be assumed that the main

canons had been excavated by glacial action more deeply than the
lateral branches, owing to the greater eroding power of the glaciers
which occupied them. This is a simple and natural explanation of
the conditions observ^ed, and if we admit the great amount of erosion
usually assumed for ancient glaciers, it must be accepted as an ade-
<juate cause for the great strength of the main channels of ice dis-
charge. To the writer it appears that the main work of sculp-
turing in the Sierra Nevada is to be attributed to water erosion,

while only minor features .... are to be referred to glacial action.
With this conclusion in mind, the great inequality in the depth of
the main glacial troughs and of their lateral branches, is too great a
work to be ascribed to the erosive power of ice" (*89, 351-352).

The hanging valleys are therefore left without explicit explana-
tion ; but it appears from other pages of the report that several of
the deeper canyons, such as Lundy, now head to the west of the
general line of mountain crest, and it is therefore possible that they
are examples of retrogressive erosion, both by water and by ice, since
the elevation of the Sierra Nevada. If this be the case, the hanging
valleys may be remnants of an ancient west^flowing drainage system,
now diverted to a more rapid eastward descent. Some such mean-
ing may be behind Russell's words : " Many of the valleys of the
Sierra Nevada .... are in fact relics of a drainage system which ante-
dates the existence of the Sierra as a prominent mountain range "
('89,348,350).

Wallace on Glaciated Valleys, 1893, — One of the most appre-
ciative statements that I have found concerning hanging valleys is
an article by Wallace on "The Ice Age and its Work," which
presents many arguments in favor of the strong erosive power of
glaciers. Wallace says : " It is evident that ice erosion to some
extent must have taken place along the whole length of the glacier's
course, and that in many cases the result might be simply to deepen
the valley all along, not quite equally, perhaps, but with no such
extreme differences as to produce a lake basin." Then after giving
much emphasis to the excavation of lake basins near the lower end
of a large glacier, where the erosive power is deductively argued to

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be at its maximum, Wallace examines several lakes to discover if
those of glacial origin have not some distinctive feature by which
they can be recognized. He points out that greater length than
breadth, and simplicity of outline, are highly significant of glacial
erosion, and that the absence of lateral bays and branches is strongly
against the theory of warping or submergence. In this connection
it is recognized that " the lake surface, not the lake bottom, repre-
sents approximately the level of the preglacial valley, and that the
lateral streams and torrents enter the lake in the way they do
because they could only erode their channels down to the level of

the old valley before the ice overwhelmed it In connection

with this subject may be noticed the many cases in which Alpine
valleys present indications of having been greatly deepened by
glacial erosion, although, owing either to the slope of the ground or
to the uniformity of the ice-action, no lake has been produced. In
some valleys, as in that of Lauterbrunnen, the trough between the
veitical rock-walls was probably partly formed before the ice age,
but was greatly deepened by glacial erosion, the result being that
the tributary streams have not since had time to excavate ravines
of equal depth with the main valley, and therefore form a series of
cascades over the lateral precipices, of which the Staubbach is the
finest example. In many other cases, however, the side streams
have cut wonderfully narrow gorges by which they enter the main
valley" ('93,754,768).

McGee's Second Paper on Glacial Canyons^ 1894,, — McGee has
given a fuller statement of the action of glacial erosion in produc-
ing discordance between lateral and main valleys in a second essay,
again entitled '* Glacial Canons," published eleven years after his
first essay on this subject. After a discussion of glacial erosion
in general, it is stated that " Glacial canons are characterized by
several peculiar features: 1. They are U-shaped rather than V-
shaped in cross-profile ; 2. Small tributary gorges usually enter at
levels considerably above the canon-bottoms; 3. In longitudinal
profile the canon-bottoms are irregularly terraced, — i. e., made up of
a series of rude steps of variable form and dimensions, — and some of
the terraces are so deeply excavated as to form rock- basins occupied

by lakelets In a region of rapid corrasion then, the main

[water] stream must .... more rapidly corrade its channel than does
its minor tributary ; and the tributary caiion must accordingly enter
its principal over a rapid or at least a convex curve in longitudinal
profile. If now the main canon become filled with ice and be

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DAVIS: GLACIAL EROSION. 315

transformed from the V to the U t^pe by its action, the distal
extremity of the tributary will be cut off and the original stream-
formed declivity replaced by the precipitous side-wall of the normal
glacier valley" ('94, 851, 359). It is explicitly stated by McGee
that this explanation does not demand great glacial erosion, because
the U-canyon of glacial origin need not be much deeper, although
significantly wider, than the preglacial V-canyon of river origin.
But the last of the above quotations postulates a special condition —
a region of rapid preglacial corrasion by streams, and in so far
does not seem applicable to the case of the Ticino or of the many
other Alpine valleys; for the well-opened slopes of the lateral
valleys, and the still wider flare on the upper slopes of the main
valleys in the Alps, provea that during their formation the main
stream must have attained a graded slope which the lateral streams
must have joined in accordant fashion; and there is nothing to
show that the open and graded floor of the main valley was sig-
nificantly trenched by river action in preglacial time. On the con-
trary, the shallowness of the trenches now found in the lateral
hanging valleys proves that even if the main valley had been
trenched, it could not have been cut down very deep.

Tarr on Cayuga Lake^ 1894, — A significant instance of discord-
ance has been pointed out by Tarr and taken by him as direct evi-
dence of the glacial erosion of a lake basin. He shows that the
north and south trough of Cayuga Lake, New York, lying in the
line of ice motion, is about three hundred feet deeper than the floor
of Salmon Creek, a tributary whose course is oblique to the ice
motion ; and he ascribes the break of grade between the two val-
leys to greater erosion in the deeper one. He generalizes so far as
to refer to Lake Ontario as probably exhibiting further instances of
discordant valleys ('9*).

De Lapparent on Hanging Valleys, 1896, — A clear and brief
statement is made by de Lapparent in his ** Le9ons de G^ographie
Physique," as if the matter were well known and undisputed.
Under the heading, " Caract^res des valines glaciaires," he writes
in effect as follows : " When a glacier disappears, the lateral val-
leys, which had been eroded before the glacial period with relation
to the local baselevel determined by the river that the glacier after-
wards replaced, may, on the disappearance of the ice, no longer pre-
sent accordant junctions with the main valley. Cascades and rapids
will therefore occur at their mouths in greater number than in a dis-
trict of the same strength of relief which has not been glaciated.

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All these featureB are dearly seen in Norway." ('96, 210; '96,
219).

Michter on Noncayy 1896, — The essay by Richter already
referred to contains a large number of excellent observations.
Regarding our special subject, he states that many side valleys in
Norway mouth high on the fiord walls, a^ if cut off in the deeper
erosion of the main valleys ; a similar relation being known in the
Alps, but of less distinctness. The discordance of valley depth in
Norway is thought to depend on the faster erosion of the main val-
leys by water or ice or both, when the side valleys and the uplands
were occupied by slow-moving n6v6. The side streams descending
from the floors of their hanging valleys have not yet cut even nar-
row clefts in the rock walls of the main .valleys ('96, 177-179).

J, Geikie on Glacial Erosion, 1898. — The recent volume on
" Earth Sculpture " by James Geikie gives a generally available access
to the results of Richter's observations on Norway. The following
quotation comes after a description of the rock walls of the fiords :
" Numerous tributary waters, some of which are hardly less impor-
tant than the head-stream, do indeed pour into the fiord, but they
have not yet eroded for themselves deep trenches. After winding
through the plateau-land in broad and shallow valleys their rela-
tively gentle course is suddenly interrupted, and they at once cas-
cade down the precipitous rock-walls to the sea. The side-valleys
that open upon a fiord are thus truncated by the steep mountain-
wall as abruptly, Dr. Richter remarks, as if they had been cut across
with a knife

*' If we admit that a fiord is simply a partially drowned land-valley,
and that the profound hollow in which it lies has been eroded by
river action, how is it that the side streams have succeeded in doing
so little work ? Why should the erosion of the main or fiord-valleys
be 80 immeasurably in advance of that of the lateral valleys?
Obviously there must have been a time when the process of valley-
formation proceeded more raj)idly along the lines of the present
fiords and their head- valleys than in the side-valleys which oj>en
upon these from the fjelds. At that time the work of rain and
running water could not have been carried on equally over the whole
land, otherwise we should find now a completely developed hydro-
graphic system — not a plateau intersected by profound chasms, but

an undulating mountain-land with its regular valleys According

to Dr. Richter, the remarkable contrast between the deep valleys of
the fiords and the shallow side-valleys that open upon them from

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DAVIS: GLACIAL EROSION. 317

the f jelds — the profound erosion in the former, and the arrest of
erosion on the plateau — admits of only one explanation. While
rivers and rapid icenstreams, flowing in previously excavated valleys,
were actively engaged in deepening these, the adjacent f jelds were

buried under sheets of n4v4 In short, while rivers and glaciers

were deepening the great valleys and making their walls steeper,
the intervening mountain-hfeights were gradually being reduced and

levelled by denudation It was somewhat otherwise in the Alps,

where the hydrographic system, perfectly regular in preglacial
times, was only slightly modified by subsequent glacial action. Yet
even there erosion proceeded most rapidly along the chief lines of
ice-flow. Were the great rock-basins of the principal Alpine valleys
pumped dry we should find the mouths or openings of the side
valleys abruptly truncated, and their waters cascading suddenly
into the ice-deepened main- valleys. For, as Dr. Wallace has shown,
it is the present lake-surfacey not the lake-^oWom, that represents
approximately the level of the preglacial valley. In a word, erosion
proceeded most actively in the main valleys, the bottoms of which
have been lowered for several hundred feet below the bottoms of the
side-valleys. Precisely the same phenomena are repeated in Scot-
land. Were all the water to disappear from the Highland lakes and
sea-lochs, we should find waterfalls and cascades at the mouth of
every lateral stream and torrent " ('98, 246-249).

It is evident from these extracts that the deepening of valleys is
regarded as greatest where lake basins have been eroded beneath the
preglacial valley floors ; and this belief is explicitly expressed in the
following extract from the latest edition of the same author's " Great
Ice Age," the standard work on that subject : '* Take the case of a
glacier creeping down an Alpine valley and spreading itself out
upon the low ground at the foot of the mountains. Let us suppose
that, in the upper part of its course, the incline down which it moves
is greater than the slope of the lower reaches of the valley. When
the glacier attains the more gently inclined part of its course, it is
evident^that its flow must be retarded, and there will therefore be a
tendency in the ice to accumulate or heap uj). Now we know that
the pressure of a body in motion upon any given surface varies with
the degree at which that surface is inclined ; as the inclination
decreases the pressure increases. It follows from this that when the
glacier leaves the steeper part of its course, and begins to creep
down the gentler slope beyond, it will press with greater force upon
its rocky bed, and this increased pressure will be further intensified

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by the greater thickness of the accumulated ice The result of

all this is the formation of a rock-basin, the deeper portion of which
lies towards the upper end, just where the grinding force of the
glacier is greatest" ('95, 228, 229).

It seems to me that too great emphasis is here placed on the ero-
sion accomplished near the end of a glacier, as indicated by lakes,
and not enough upon the deepening of the valleys up-stream from
terminal lakes, as indicated by hanging valleys. It is also to be
noted that de Lapparent, Richter, and J. Greikie all describe the
hanging valleys of Norway as if their preglacial form had not been
significantly changed, thus failing to bring clearly forward the fact
that the valleys of today are the ice channels of the past, and that
the larger and smaller channels must have normally discordant floors
in a system of glacial drainage, just as they have in a system of river
drainage, although to a much greater degree. The full analogy
between ice and water channels, which throws so much light on the
whole question of glacial erosion, was first clearly set forth by the
two following observers.

Gannett on Lake Chelan^ 1898, — The most complete statement
of the general principles involved in the production of hanging
valleys that I have found in print is in an article on Lake Chelan, in
the Cascade Mountains of Washington, by Henry Gannett. Chelan
is a long narrow lake occupying the distal two thirds of the deep U-
shaped valley of the Stehekin River on the eastern slope of the
mountains. It was occupied in the glacial period by a heavy ice
stream, fifty or sixty miles long, and half a mile to a mile broad.
The rock walls which enclose the valley are strikingly parallel to
one another, without buttressing spurs ; they rise 4000 to 5000 feet
above the lake waters. Nearly all the streams which flow into the
valley tumble over its walls in a series of cascades. " From all
indications it appears that the ice must have been at least 3000 feet
deep in this gorge of the Stehekin, since several of the smaller
branches joinr the main glacier at that height above its bed.'*

Speaking of these features in a more general way, Gannett says :
" A glacier is a river of ice, and it behaves almost precisely as a
river of water does. Its effects upon its channel are almost pre-
cisely similar to those of a river upon its channel, excepting in the

fact that all its operations are on a vastly greater scale A word

of caution must here be interpolated. The channel of a river, in
wliich its water flows, must not be confused with its valley, which
it drains. ^Phe above comparison refers to the channel of a river

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[or of a glacier], not to its valley The glacier moves down the

gorge, scouring and cutting the bottom and sides as it travels.
The ends of the mountain spurs are planed off instead of being
trimmed to sharp, angular points, as is done by streams in gorges

cut by them Where the main glacier is joined by a branch, the

bed of the branch is commonly found to be at a higher level than
the bed of the main glacier, because being larger and heavder the
main glacier has greater cutting power ; indeed, in many cases the
beds of small branches are hundreds, or even thousands, of feet
higher than that of the main glacier to which they are tributary.
The parallelism between the glacier and the river in their channels
is further illustrated by this fact. The surface of the ice in the
main glacier and in the branch must have been at the same level,
although the bottoms, as stated above, differ greatly in elevation.
So it is with a river at the point of junction of branches. The
surface of the water must be practically at the same level in all
cases, but the bottoms of the channels differ by the difference in
depth of the streams at their point of junction. This fact affords
us a measure of the minimum thickness of the ice at any place. It
cannot have been less than the vertical distance between the bed
of the main glacier and that of the tributary, and, indeed, must in
all cases have been greater, owing to the thickness of the tribu-
tary" ('98, 417-428, especially 418-420).

Penck on Alpine VcUleya^ 1899, — A no less explicit and detailed
statement of the peculiar features of glacial channels and their rela-
tions to river channels was made by Penck at the meeting of the
International Greographical Congress in Berlin, September, 1899.
The discordance of lateral and main valley floors wa« described as a
general feature of all the larger Alpine valleys within their glaciated
areas. The possibility of explaining the discordance by faulting, as
suggested by Rothpletz for the Linththal, was considered, but rejected.
The contrasts of the glaciated and non-glaciated Alpine valleys were
strongly emphasized. The excess of the depth in the main valley
beneath the floor of the hanging ^laterals was taken as a minimum
measure for glacial erosion, and the term " over-deepened," already
adopted on earlier pages of this essay, was applied to valleys thus
worn to a greater depth than would have been possible to normal
rivers. The publication of Penck's address is awaited with interest.

Harker on Glacial Valleys in Skye, 1899. — A brief article by
Harker on glaciation in Skye describes the valleys as eroded in
massive gabbros, with U-shaped cross-section, especially in the upper

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stretches, and frequently heading in a corrie whose floor may hold a
small rock-basin tarn. In longitudinal profile, the floor of a valley
often consists of two or three stretches of relatively gentle slopes,
or sometimes of basin form and then holding lakes, separated by
relatively sudden descents. Tributary valleys mouth at a consid-
erably higher level than the floor of the main valley, (*99, 196-199).

Gilbert on Alaskan Valleys^ 1899. — A valuable contribution to
the origin of hanging valleys will be found in a report on the Harri-
man Alaskan Expedition of 1899, to be published shortly. A gen-
eral statement of results wa« made by Gilbert during the session of the
Geological Society of America in Washington, December, 1899, when
the importance of the hanging lateral valleys in the Alaskan fiords,
and their bearing on the problem of glacial erosion, was clearly set
forth.

Blanford on Scotch GlenSy 1900. — The only article that I have
found on hanging valleys in Scotland is by Blanford, " On a partic-
idar form of surface, apparently the result of glacial erosion, seen on
Loch Lochy and elsewhere." The '* particular form " here referred
to is the smoothness of the sides of the Great Glen of Scotland, a
feature that may be held analogous to the smooth rock walls of the
Norwegian fiords, and to the spurless basal cliffs of the glaciated
Alpine valleys. It is inferred that in preglacial time the streams of
the lateral glens were separated by advancing spurs which buttressed
the sides of the Great Glen. Now the spurs seem to have been
truncated, producing the smooth and even sides of the glen, to
which attention is esj>ecially directed. The lateral glens are described
as at present opening a thousand feet above the floor of the Great
Glen, whose smoothed sides are very little eroded by the descending
tributary streams. The change from the inferred preglacial form is
conservatively taken to indicate a glacial erosion of "at least 250 or
300 feet of rock " (1900, 198-204).

Hershey on Sierra Costa^ California^ 1900. An article by Her-
shev, already referred to above, is the latest contribution to the sub-
ject in hand. In following uj) a valley in the Sierra Costa in
northwest California, it is at first V-shaj>ed, with jagged ledges
between sharjMJut ravines on the sides, and hardly wider at the
bottom than the stream that drains it. On reaching the stretch
once occupied by a local glacier, the valley becomes an open U-
shaped trough, with smooth slopes free from ravines and spurs.
Above the limit of glacial smoothing, the mountain sides are still
deeply scored with ravines and jagged with outcropping ledges.

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The descent of a glaciated valley floor is effected by a series of
steps ; the stretches of more gentle fall alternate with almost pre-
cipitous falls where the floor is let down several hundred feet,
Corries with tarns in their floors are well developed (1900, 42-57) ^

Several essays by Norwegian authors remain to be considered. It
has not been possible to make reference to them without postponing
the appearance of this paper, and consideration of them is there-
fore deferred to another occasion.

With all these new contributions to the subject, it may be
expected that hanging lateral valleys and overdeepened main valleys
will soon gain the importance that they deserve in geographical
literature.

LITERATURE.

Barrett, R. L.
1900. The Sundal drainage system in central Norway. Bull. Amer. geogr.
soc, 32, (in press).
Blanford, W. T.
1900. On a particular form of surface, apparently the result of glacial ero-
sion, seen on Loch Lochy and elsewhere. Quart journ. geol. soc, 66, pp.
198-204.
Boule, M.

'96. La topographic glaciaire en Auvergne. Ann. de g^ogr., 5, pp. 277-
296.
BrUckner, E.
'85. Die Vergletscherung des Salzachgebietes. Geogr. Abhandlungen,
Vienna, 1, pp. 1-188.
Davis, W. M.

1900. Glacial erosion in the valley of the Ticino. Appalachia, 9, pp. 136-156.
Forbes, J. D.

'53. Norway and its glaciers. Edinburgh.
Forel, F. A.

*97. Fleuves et glaciers. Bull. soc. vaud. sci. nat., 33, pp. 202-204.
Gannett, H.

'98. Lake Chelan. Nat. geogr. mag., 9, pp. 417-428*
Geikie, J.
'95. The great ice age. New York. 3d ed.
'98. Earth sculpture. London.

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322 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

Hall, J.

*43. Geology of New York. R. 4, comprising the geology of the fourth dis-
trict.
Harker, A.

'99. Glaciated valleys in the Cuillins, Skye. Geol. mag., 6, pp. 196-199.
Helm, A.

'78. Mechanifimus der Gebirgsblldung. BaseL

'79. Ueber die Erosion im Gebiete der Reuss. Jahrb. echw. Alpenclab,
14, pp. 371-405.
Hershey, O. H.

1900. Ancient alpine glaciers of the Sierra Costa Mountains in California.
Journ. geol., 8, pp. 42-67.
Lapparent, A. de.

*96, *98. Le9on8 de geographic physique. Paris. 1st ed., 1896 ; 2d ed., 1898.
Lubbock, J.

*96. The scenery of Switzerland. New York.
Lug6on, M.

'97. t«e9on d'ouverture du cours de gfeographie physique profess^ a I'uni-
ver8it6 de Lausanne. Bull. soc. vaud. sci. nat., 33, pp. 49-78.
McGee, W. J.

'83. Glacial canyons. Proc. Amer. assoc, 1883, p. 238.

'94. Glacial canyons. Journ. geol., 2, pp. 350-364.
Marr, J. E.

1900. The scientific study of scenery. London.
Meunier, S.

'97. Sur r allure g^n^rale de la denudation glaciaire. Comptes rendus, 124.
p. 1043.
Play fair, J.

*02. Illustrations of the Huttonian theory of the earth. Edinburgh.
Richter, E.

'96. Geomorphologische Beobachtungen aus Norwegen. Sitzungsber. k. k.
Akad. Wien, math, naturw. Classe, 105, Abth. 1, pp. 147-189.
Rothpletz, A.

'98. Das geolektonische Problem der Glamer Alpen. Jena.
Russell, I. C.

'89. Quaternary History of Mono Valley, California. 8th ann. rep. U. S.
geol. surv., 1889, pp. 261-394.
RUtimeyer L.

'69. Ueber Thai- und See-Blldung. Basel.
Tarr, R. S.

*94. Lake Cayuga a rock basin. Bull. geol. soc. Amer., 5, pp. 339-356.
Wallace, A. R.

'93. The ice age and its work. Fortnightly rev., 60, pp. 616-633, 760-
774.

'96. The gorge of the Aar and its teachings. Fortnightly rev., 66, pp.
175-182.

Printed July^ 1900,

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Da VIA. — Glacial Erosion.

PLATE 1.

Fig. A. Valley of the Ticino, looking up-stream to Giomico.
Fig. B. Cliffs beneath graded slopes, eastern arm of Lake Lugano.

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.as

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i
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Da viB. — GlftcUU Erosion.

PLATE 2.

Fig. A, Cascade at tlie mouth of a hanging valley, eastern arm of Lake

Lugano.
Fig. J5. Sterling Fall, Milford Sound, New Zealand.

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X

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Da VI*. — Glacial Erosion.

PLATE 3.

Fig. A. The Strandfos, a cai^cade descending from a hanging valley to Sandven

Lake, Norway.
Fig. B. Valley of the Romanches Alps of Dauphiny.

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h
X

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AUG 8d 1901

No. 18. — The Polychaeta of the Puget /Sound Region.

By Herbert Parlin Johnson.

The following account of the Polychaeta of Puget Sound and
neighboring waters is primarily based upon a collection made by
Nathan R. Harrington, lately deceased, a member of the Columbia
University Zoological Expedition to that region in the summer of
1896. The collection, comprising thirty-four species belonging to
seventeen families, was sent to me in January, 1898. Preliminary
examination showed that the collection, although meagre, in part
poorly preserved, and almost destitute of data, contained much of
interest. It seemed best, however, to defer any publication of
results until more and better material could be obtained. Thanks
to the good efforts of several collectors, notably Prof. William E.
Ritter and Miss Alice Robertson, both of the University of Cali-
fornia, very substantial additions have been made to the original
collection. These, together with two species from Victoria, B. C,
kindly contributed by Prof. William A. Herdman, raise the total
number of species to fifty-one, distributed in thirty-four genera and
tw^enty-six families (see Table, p. 384) — practically all the Poly-
chaeta known to occur in the Puget Sound region.^ They cannot
reasonably be supposed to rej)re8ent more than a fourth or even a
fifth of the actual Polj'chaete fauna. Nearly all the collecting thus
far has been between tide-marks. Dredging has been small in
amount and limited to very moderate depths (not over thirty fath-
oms) ; and the entire absence of pelagic forms — with the possible
exception of Arichhopsis megnlops — would indicate but slight use
of the tow-net.

1 Ehlers ('68) describes eight species from the Pacific Coast ; live of these [Nereis
agasfizi, N. procera, JV. vejrillosa, X. virens and XepMhys coeea) have been found in
Puget Sound or vicinity.

Balrd ('63) describes nine species, all from Esquimalt Harbor, Vancouver- Island.
Four of these. Polynoe ( Lepldonohui) itisignls, lordi, frur/Uia, and XerelsfoHata (= X.
virens Sars) have been identified, and an account is given of them. Lepidonotxis
gntbei Is in all probability the same as Pohjtioe itinhjiiUs. There remain unidentified
only four of Baird's species (Hannot/ute unicolor, Xereis hicamilicufata, Glycera cor-
nigata, &nd Sabellaria aaxicara^ These it is practically impohsible lo identify with
certainty from Baird's descriptions.

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582 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

Although comparison has been carefully made with the descrip-
tions of Japanese Polychaeta given by Grube ('77) and by von
Marenzeller (79 and '84), no species, with the sole exception of
the widely ranging Ilarmothoe imbricata, has been found to
inhabit both sides of the Pacific. Thb is not surprising when we
consider that nearly all of the sixty-two species of Grube's and
Marenzeller's lists are from the southern portions of Japan, and
therefore belong to the Indo-Pacific fauna, not to the circumboreaL
But, on the other hand, comparison of Puget Sound Annelids wi^
those of Bering Sea (Wir^*n, '83; Marenzeller, '90) also shows
absolute dissimilarity. So far as known, the Polychaeta of the
more northern parts of Bering Sea — the only ones of that region
dealt with by Wir^^n and von Marenzeller — are practically those of
the Arctic and North Atlantic (i. e., are circumboreal) with very little
^mixture of forms peculiar to the North Pacific. In Wir^n's list
-of twenty-nine species there is not one which can be regarded as
ibelonging distinctively to the North Pacific. Yon Marenzeller's
list of twenty-four (exclusive of eight previously given by Wiren)
affords only two new species; the others are well-known inhabi-
tants of the shores of Scandinavia, Iceland, Greenland, or north-
<eastern America. The data as yet available are wholly insufli-
cient to justify even a rough estimate of what proportion of the
Puget Sound Annelids range far to the northward and eastward
along the Aleutian chain. From such a category should of course
be excluded purely circumboreal and North- Atlantic species (e. ^.,
jPoli/?ioe {Le2)ido7iotus) sqiunnata, Ilarmothoe imbricata^ JSereis
virens), as these are no more characteristic of the Pacific than of
the Atlantic. Wlien the extensive series of Polychaeta collected
by Professor Ritter and Dr. Coe during the Harriman Alaskan
Expedition of 1899 shall have been worked up it will doubtless be
found to contain many species which range southward to Puget
Sound and beyond.

Descriptions of localities where collecting was done by the
Columbia Expedition have already been given in the general reports
by members of the Expedition ('97, '97a) ; and the topography of
the region, with indication of collecting stations, is shown in a map
published with the first of these reports. In many instances the
authors mention the occurrence of Annelids ; but it is not always
possible to determine accurately what species is meant. Moreover,
several species are mentioned — by generic name at least — that
are not represented in the collections.

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JOHNSON; POLYCHAETA OF PUGET SOUND REGION. 383

In the summers of '97, '98, and *99 Miss Robertson collected in
the vicinity of Seattle, a region much further up the Sound than
that investigated by the Columbia University Expedition. About
Seattle the shores are generally beaches of muddy sand, but some-
times clean sand, as is the case between Duwanish Head and Alki
Point. Alki Point itself is rocky, and in 1898 numerous Annelids
were found between and under the stones. Under the bluff on the
north shore of Port Orchard Channel (stretching westward from the
Sound) extends Pleasant Beach, composed of muddy sand. Numer-
ous burrowing forms are from this locality. On this beach Professor
Ritter collected several species of Polychaeta in 1899.

Following is a list of the species represented in the various
collections, which shows, so far as known, the local and the
geographical distribution of each species. For convenience' sake,
the two districts explored by the Columbia University Expedition are
designated " Port Townsend Region " (including Sequim Bay, Dis-
covery Bay, Scow Bay, and Hood's Canal), and " Neah Bay " (at the
entrance of Straits of Juan de Fuca) ; while that portion of Puget
Sound investigated by Miss Robertson and Professor Ritter is indi-
cated as " Seattle Region " and comprises not only the immediate
vicinity of Seattle but also Port Orchard Channel and its shores —
Orchard Point, Mud Bay, "Port Orchard," Pleasant Beach, and
Channel Rocks. A + indicates occurrence of a species in any given
region.

Two species {N'orthia iridesce?*^ sp. nov. and Sterndsjns fossor
(?) Stimpson) were dredged by Prof. W. A. Herdman at Victoria,
Vancouver Island, B. C.

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384 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

G«ogri4>hical Distribution.

2.
3.

4.
5.

10.

11.

12.

13.

14.
15.
1«.
17.

18.

10.

20.
21.

22.
23.

24.

25.
2(1
27.

28.

POLYKOIDAE.

Polynoe (Lepidonotus) squamata

(L.) Aud. and M. -Edwards.
P. insi^U (Baird)
P. lordi (Baird)

P. pulchra Johnson

P. fragilis (Baird)

Harmothoe imbricata (L.) Malm-

gren.
H. ipiiionelloides sp. nov.
H. complanata sp. nov.
H. paciflca sp. nov.
H. tuta (Grube)

SiGALIONIDAE.

Sthenelais fiisca Johnson

Hesionidak.
Podarke pugetteiisis sp. nov.

NivKEIDAE.

Nereis virens M. Sars

N. vexillo.sa Grube
N. agassizi Khlers
X. cyclurus Harrington
N. procera Khlers

Nefhthvdidae.
Xephthys coeca (Fab.) Oerst.

EuPHROi*YNn>AE.

E uphrosyne heterobraiichia sp
nov.

Syllidae.
Piouosyllis elonaata sp. nov.
TiypaiKKsyllLs gemmipara sp.'
nov. !

Onupiiididae. ,

Northia elegans sp. nov. ;

N. iridescens s]). nov. j

LCMBinCONEREIDAE. |

Liunbricrmert'is zonata sp. nov.,
Glycerii>ae. I

Glycera riigosa sp. nov.
G. nana s]). nov.
Heniipo<lia boreal is sp. nov.

Aricudae.
Scoloplos elongata sp. nov.

+ 4-
4-

H-

Circumboreal ; Cal. coast to Santa

Monica.
Bering Sea to San Diego.
Northward (?); Cal. coast to

San Francisco.
Cal. coast to Pacific Grove,
** " " San Francisco.
Circumboreal; Japan; Cal.

coast.
Northward.
Cal. coast (San Diego).

Northward ; Sitka (Grube).

Cal. coast to San Pedro.

Cal. coast (Pacific Grove).

Northern Europe & N. E. coast
of N. A. ; Cal. coast to Bolinas.
Bering Sea to Santa Barbara.
Southward to Santa Barbara.

Gulf of Georgia (Ehlers).

Circumboreal ; Alaska to Toma-
les Bay, Cal.

Southward to Monterey Bay.

Victoria, B. C. (Herdman).

4-' + !
+ I

+ 1 Southward ; Cal. coast to Toma-
i les Bay.

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JOHNSON: POLYCHAETA OF PUGET SOUND REGION. 385

1
1

G«ogtAphlcaL Dl^trlbutLoD,

Lkvisseniidae.

29.

Arlcideopsm megalopa g&iL et sp.

■

L -

IIOV*

MAc*eumit>Jii£.

80.

Magelonn lon^corQiJ^ Ap. iiov.

Capitelmc^ae.

Capital h dizoiiata *p. nov.

Chluha1vMII>ai:.

82.

Trophonia papillata sp. noy.

Southward; Shelter Cove, CaL

SS.

FlabelUgera infundibularis sp.

nov.

Stbrkaspidab.

d4.

Sterxuwpis foflsor (?) Stimp.

Victoria, B. C. (Herdman) ; Ber-
ing Sea, Japan (Maren-
zeUer); N. E. coast North

America.

Maldanidas.

86.

Clymenella rubrocincta sp. noy.

Southward to San Pedro, Cal.

86.

Nicomache pereonata sp. noy.

Ammocharidab.

87.

Ammochares occidentalU sp. noy.

Arbnicolidae.

88.

Mediterranean.

CiRRATCLIDAB.

89.

Cirratulua cingulatus sp. nov.

40.

C. robustus sp. nov.

Amphictenidae.

41.

Pectinarla brevicoma sp. nov.

Ampharetidab.

42.

Sabellides anops sp. nov.

Terebellidae.

48.

Amphitrite robusta sp. nov.

44.

A. spiralis sp. nov.

45.

Lanice heterobranchia sp. nov.

46.

Thelepus crispus sp. nov.

Southward to San Francisco.

Sabellidae.

47.

Bisplra polymorpha sp. nov.

Southward to Monterey Bay.

48.

Megachone aurantiaca gen. et.

-f

sp. nov.

Eriographidae.

40.

Myxicola pacifica sp. nov.

H-

Seupulidae.

60.

Serpula Columbiana sp. nov.

Southward to San Francisco.

61.

Serpula zygophom sp. nov.

Northward (?).

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386 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

The entire absence of Phyllodocidae and Opheliidae from a col-
lection of this size and representative character is remarkable. The
Eunicidae and Chaetopteridae are represented, the former by several
headless fragments of some large species, and the latter by numerous
long, annulated, empty tubes.

For any zoologist devoted to the study of living animals, the
examination and description of a collection of preserved spedmens,
gathered by others in a region he has never visited, is likely to
prove an irksome and unprofitable task. In the present instance,
however, the task has been enlivened by the discovery of several
very interesting forms. First and foremost should be mentioned
Tri/proiosi/llia gemmipara, a Syllidian with alternation of genera-
tions in which the sexual zooids, instead of forming a linear series,
arise by collateral budding near the posterior extremity. Hamio-
thoe tufa, with its great number of asymmetrical somites, also
deserves notice ; and as regards zoogeography, the discovery of a
Pacific species of Mai/elona, the finding of a genuine Hemipodia in
the Northern Hemisphere, and the confirmation of Gamble and
Ash worth's (^00) statement regarding the occurrence of Arenicola
claparedei on the west coast of America, are worthy of note.

I gladly avail myself of this opportunity to express my sincere and
hearty thanks to Prof. C. O. Whitman, who generously placed at
my disposal an investigator's room at the Marine Biological Labora-
tory during the latter part of the summer of 1900 ; to Prof. E. L.
Mark, of Harvard, to whom I am indebted for laboratory privileges
at the Museum of Comparative Zoology; and to Dr. Wm. M.
Woodworth for permission to make use of the Polychacte collec-
tions of the same institution.

POLTNOIDAE.

1. Polynoe squamata (L.) Aud. and M.-Edwards.

Lepidonotics squamatus Leach. Zoological miscellany, London,
1816.

This well-known circumboreal species is represented by three
specimens, probably from the vicinity of Port Townsend.

P. squamata occurs on the California coast as far south as Santa
Monica, where a specimen was obtained for me by Mr. J. J. Rivers,
the well-known entomologist. At Pacific Grove it is frequent in

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JOHNSON : POLYCHAETA OF PUGET SOUND REGION. 387

dredgings at twelve fathoms and over, but T have never obtained it
there between tide-marks. I have collected it just above low-water
mark at Point Cavallo, on the northern shore of the Golden Gate,
and even higher on the beach near Black Point, Sonoma County,
about one hundred miles north of San Francisco. The specimens
from Puget Sound are larger than any as yet collected from the
California coast, but are inferior in size to New England specimens.
It is probable that this species will be found to have a wide distri-
bution along the eastern and western shores of the North Pacific,
comparable to its dispersal on both sides of the North Atlantic.

2. PolynoeinsigxiiB (Baird).

Xf€pidonotii8 insignis Baird. Proc. zool. soc. London, Apr.,
1863, p. 106.

II(do8ydna insignis Baird. Joum. Linn. soc. London, vol. 8.
(Zoology), 1865, p. 188.

Polynoe brerisetosa (Kinberg) Johnson. Proc. Cal. acad. sci-
ences, 3d ser., Zoology, vol. 1, 1897, p. 167. Figs. 24, 31, 40, 40a,
46, 46a.

This, the commonest Polynoid of the western coast of North
America, is represented in the Puget Sound collections by several
slender, darkly pigmented specimens commensal with Thelepus
crispuSy and a single specimen with remarkably thick and tuber-
culated elytra, obtained by the Columbia University Expedition;
also by four specimens collected by Miss Robertson at Alki Point.

With the exception of Ilarmothoe imbricata this species has the
widest known distribution along our western sea-board of any of its
family, ranging from San Diego to Kadiak.^ South of Point Con-
cepcion it is rare, at least inshore, being almost wholly replaced by
P. ccUi/ornica,^ Its remarkable variations according to habitat have
been described elsewhere (vide Johnson, *97, p. 167).

The identification of this species with the Halosydna brevisetosa
of Kinberg ('55; '58, p. 18), as given by me in the "Preliminary
Account" ('97), was undoubtedly characterized by too little confi-
dence in the accuracy of Kinberg's figures and too much influenced
by his statement that Haloaydna bremsetoaa was collected in San-
(salito Bay, near San Francisco. As P, insig^iis is the only Poly-

1 A smaU collection of Polychaeta from Kadiak Island, including a single P. insignU^
wai gathered by Mr. Cloudsley Rutter, and kindly loaned to me for examination and de-
scription by Stanford University.

s Name changed from P. reticulata Johnson C97) as the latter name is preoccupied.

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388 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

noid with eighteen pairs of elytra I have ever seen from that
portion of the coast, it is highly probable that Kinberg's species
came from another part of the world. P. insignia was described
by Baird in 1863 from specimens collected by J. K. Lord at
Esquimalt, Vancouver Island. His Lejndonotiis grubei^ described
at the same time (and subsequently) with the foregoing, is in all
probability a mere color variety of P. insignia,

8. Polynoe lordi (Baird).

Lepidon(Ptns lordi Baird. Proc. zool. soc. London, Apr., 1863.

Halosydna lordi Baird. Journ. Linn. soc. Ldndon, vol. 8 (Zool-
ogy), 1865, p. 190.

Polynoe lordi Johnson. Proc. Cal. acad. sciences, 8d ser., Zool-
ogy, vol. 1, 1897, p. 175. Figs. 35, 44, 61.

As in the case of P. insignia and P. fragilis^ the Paget Sound
region may be regarded as the type locality of the present species.
All three were collected by J. K. Lord at Esquimalt, Vancouver
Island. For an instructive and entertaining account of this curious
Polynoid's habits and habitat, the reader is referred to Lord's
<< Naturalist in Vancouver Island and British Columbia,'' vol. 2,
page 9 ('66). As this somewhat rare work is doubtless inaccessible
to many, I quote somewhat at length from Lord's account. Speak-
ing of P'iaaurella cratitia^ the host of P, lordi, he says: "I had
found him at last and at home, so pounced upon him as a law-
ful and legitimate prize. Knife and hammer soon severed his
close attachment to the rocks ; and turning him up, to take a peep at
his powerful ring of muscle and strangely-formed breathing appa-
ratus, I spied a worm evidently very uneasy, about three inches long,
brown, and in shape like an ancient dagger blade. He appeared to
me to be wriggling out from betwixt the folds of the foot or the
mantle, and apparently most anxious to escape .... In displacing
other shells, I found in nearly every one a similar tenant: the
secret was discovered, the worm was a parasite, that lived in
peace and good-fellowship with the Keyhole. . . . That the parasite
worm does no harm is clearly proved by the healthy state of the
mollusc in whose shell it takes up its abode .... On more carefully
examining the position of the worm I found it was invariably
coiled away in a semi-circle under the foot, like a ribbon on its edge,
never fiat. This seems to me a wise provision ; for the pressure of
the muscles when the limpet grips the rock would crush a soft-bodied
worm to death, if flat ; but by being edge on, which is the position

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JOHNSON: POLYCHAETA OF PUGET SOUND REGION. 389

chosen, all risk of harm is avoided, as it fits in a cleft between two
layers of soft material. ... At least four out of every six contained
a parasite, and, what is rather strange, the worms ^-ere nearly all
of one size."

Later collectors have found it not uncommon in that region. The
Columbia University collection contains four specimens, probably
from Neah Bay, as the species is mentioned by Harrington and Grif-
fin ('97) as having been found there "on JFissurdlay Miss
Robertson's collections afford only a single fragmentary specimen
from Alki Point. I have several from Anacortes, collected by Miss
Louise M. Carpenter, of Berkeley, Cal. These last occurred under
the mantle of specimens of Olyphia ctapera, pre8er> ed with the Anne-
lids. This Gastropod is undoubtedly the usual host, but I have
found it also in the gill-groove of CryptorMton ateUeri, and once on
the Leather Star (Dermasterias imbricata), crawling on the aboral
surface. The southern limit, so far as known, is Point San Pedro,
about twelve miles south of San Francisco. Two specimens were
found at that locality by Prof. W. J. Raymond of the University
of California, who kindly placed them at my disposal. Although
about eighteen specimens of Olyphis aspera were examined, only
these two specimens of P. lordi were obtained. Like Pdynoe
fragilis^ this spedes becomes more abundant northward. Every
specimen of Olyphis cupera brought from Anacortes by Miss
Carpenter had one or two examples of P. lordi under its mantle ;
and Lord states the proportion to be "at least four out of every
six."

Two of the Columbia University specimens have almost no pig-
ment. Even the brown zone on the ninth somite, so constant a feature
of , this species, cannot be made out in one specimen. This example
is remarkable also for its size, having 83 somites and 41 pairs of
elytra.

P. lordi J like Lepidametria commensalis (Webster, '79), Polynoe
gigas Johnson, and Harmothoe tuta (Grube) often has asjinmet-
rical somites in the posterior portion of the body. In one of the
specimens from Anacortes the thirty-first somite is asymmetrical
(cirriferous on the right side, elytrophorous on the left). In a
specimen from Dillon's Beach, Sonoma Co., Cal., there are as many
as nine asymmetrical somites, and yet there are the same number
of elytra (27) on both sides. The elytra on the right side are borne
on somites 2, 4, 5, 7 ... . 23, 26, 28, 29, 31, 33, 35, 37, 38, 40, 42,

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390 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

44, 46, 48, 50, 52 ; and on the left side on somites 2, 4, 5 23,

26, 28, 29, 31, 33, 35, 37, 39, 41, 43, 44, 46, 47, 48, 50, The asym-
metrical somites are printed in heavier type.

4. Polynoe pulchra Johnson.

Polytioe pulchra Johnson. Proc. Cal. acad. sciences, 3d ser^
Zoology, vol. 1, 1897, p. 177. Figs. 34, 43, 43a, 50, 50a, 50b.

The species of scaly annelid referred to by Harrington and Griffin
(*97) as occurring on Molothutia califarnica is without much
doubt P. pidchra, as I have found it a frequent commensal or parasite
of that Holothnrian. It is well represented in the Columbia coUec-
tion. None of the specimens exhibits dark brown or black mark-
ings on the elytra, but all appear to have had the protective reddish
or flesh tints characteristic of the individuals found on Holothuria
califorjiica.

5. Polynoe fragilis (Baird). PI. 1, fig. 1.

Lepidonotus fragilis Baird. Proc. zool. soc. London, Apr., 1863.

IlcUoaydna fragilis Baird. Journ. Linn. soc. London, vol. 8
(Zoology), 1865, p. 190.

Polynoe fragilis Johnson. Proc. Cal. acad. sciences, 3d ser.,
Zoology, vol. 1, 1897, p. 179. Figs. 36, 45, 52, 52a, 52b.

Numerous specimens from all three districts of the Puget Sound
region. As stated in the "Preliminary Account" (Johnson, '97,
p. 180) this species is much more plentiful in the Puget Sound
region than on the California coast, where I have collected it only
in San Francisco Bay, and only three or four specimens in as many
years. Its great abundance at Port Orchard is noted by Miss
Robertson as follows : " Twenty specimens were taken from twenty-
seven or twenty-eight Star- fishes. Several times two and in one
instance three, were found on a single Star-fish."

The frequent absence of ventral cirri is a striking peculiarity.
Even when present, the ventral cirrus is very diminutive ; but, on
the other hand, its absence on all the parapodia seems to be a rare
occurrence. Of the twenty-nine examples at my command, only
one (and an imperfect specimen at that) is entirely destitute of ven-
tral cirri. In not a single individual, however, is every parapodium
provided with a ventral cirrus.

6. Harmothoe imbricata (L.) Malmgren.

This ubiquitous species was collected by the Columbia Expedi-
tion, also by Miss Robertson at Alki Point and at other places.
Most of the specimens are of the usual greenish gray tint with

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JOHNSON: POLYCHAETA OF PUGET SOUND REGION. 391

mottled, iron-gray elytra. One of the largest specimens obtained at
Alki Point has numerous undeveloped ova attached to the dorsal
setae and in the intersegmental furrows. These bunches of ova are
only partially covered by the elytra.

7. Harmothoe iphionelloides sp. nov. PI. 1, figs. 2-7.

Form short, broad, and flattened ; width, including elytra, two
fifths of length; dorsum and prostomium completely covered by
elytra, which overlap extensively; somites, 36; elytra, 15 pairs,
borne on somites 2, 4, 5, 7 23, 26, 29, 32.

Prostomium (Fig. 2) broad, ti*ansverse diameter exceeding the
longitudinal ; distinctly bilobed with median sulcus ; the lobes ovoid,
rounded in front. Palpi stout, fusiform, minutely papillated, trans-
versely ringed with grooves, acuminate. Tentacle with immense
basal joint, nearly one half its length; terminal segment slightly
papillate, subulate, unpigmented. Antennae with basal joints about
as long as tentacle, slightly bulbous near tip, with subterminal dark
band ; slightly papillated. Eyes large, black, the posterior pair
smaller and nearer together than the anterior pair,

Peristomicd cirri (Fig. 2) much stouter than antennae, and about
twice as long ; with filiform papillae near tips ; basal joints very
long; subterminal bulbous enlargement, with dark pigment-zone.

Elytra thick, with large, rough, irregularly polygonal, flattened
tubercles (Fig. 3), forming a pattern like alligator skin; tubercles
increase in size from the concave (protected) side of elytron towards
the convex and exposed portion ; elytra become larger towards
middle of series, and diminish again towards posterior end ; ciliate
on outer margin ; except first pair, all elytra more or less strongly
reniform, the concave edge of each embracing the preceding elytro-
phore.

Parapodia (Fig. 4) rather long, thick, biramous; each ramus
ending in a finger-like prolongation, into which the acicula extends.
Dorsal cirrus extends beyond the setae ; basal joint nearly one fifth
its length with pin-head-like papillae towards the incrassated tip.
Ventral cirrus subulate, slightly papillate. Ventral setae straw-
colored, hardly extending beyond the longist dorsal (Fig. 4, below
the dorsal cirrus) , and only slightly stouter than the uppermost dor-
sal (c/. Figs. 6 and 6); slightly hooked at tip, with 12-20 "frills"
of usual form. Dorsal setae white, forming a graduated series from
uppermost stout, short, strongly-curved ones (Figs. 4 and 6) to the
lower elongated slender ones (Figs. 7 a, h) . The fine serrations ex-
tend nearly one half the length of seta, whether it be long or short.

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392 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

XephricUal papMie very short, cylindrical, length and diameter
about equal ; begin at 8th and extend to 30th somite.

Length (approximately) 23 mm. ; width, including setae, 10.5 mm. ;
dorso-ventral thickness, 8.5 nmi. ; proboscis, 4 mm.

A single specimen from Pleasant Beach, near Seattle, collected
by Professor Hitter. It is chiefly notable for its great, overlapping
elytra, with tuberculation resembling the areolae of Samwtkoe
hirsuta^ or of IphioneUa.

8. Harmothoe complanata sp. nov. PI. 2, figs. 8-13.

Fortn of moderate length, flattened dorso-ventrally ; breadth, in-
cluding setae, two sevenths of length ; number of somites, 36-38 ;
elytra, 15 pairs, borne on somites 2, 4, 5, 7 23, 26, 29, 32.

Prostamium (Fig. 8) approximately six-sided, width and length
equal, deeply incised for reception of basal joint of tentacle, indis-
tinctly bilobed. Eyes four, minute, posterior pair dorsal, anterior
pair dorso-lateral, and further apart than the posterior. Tentacle
with short basal joint, which extends a little beyond the '' peaks " of
the prostomium, terminal portion nearly twice the length of the pro-
stomium. Antennae very short, inserted below level of tentacle,
two-jointed, less than one half the length of the prostomium. Palpi
very thick at base, terete, gradually and uniformly tapering to an
acute tip, barely papillate, as long as perittomial cirri

Peristomial cirri long and slender, gradually and uniformly
tapered, without subterminal dark band, slightly papillate.

Elytra (Fig. 10) thin, oval, translucent, with minute, scattered
conical tubercles.

Parnpodia (Fig. 9) long, rami distinct, finger-like tips long and
slender. Dorsal cirrus like peristomial, long and slender, extending
far beyond tips of setae, papillate. Ventral cirrus long, evenly
tapered, papillate. Dorsal setae of two sorts : (a) a supra-acicular
fascicle of very stoat, minutely serrated setae (Fig. 11) which are
the thickest in the foot ; they are arranged in a whorl, the shortest
being the uppermost and most anterior; {h) a smaller tuft of very
slender, elongated, serrulate setae (Fig. 12) inserted in the finger-
Hke process ; these are considerably longer than the preceding, and
much fewer (4-5 in number). The ventral setae (Fig. 9) are
arranged in a graduated series of which the uppermost closely re-
sembles in length, slenderness, and serrulation the dorsal set^ of
fascicle h ; the middle and lower ones are of the more usual type
(Fig. 13), with a series of "frills" near tip, beginning with very

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JOHNSON : POLYCHAETA OF PUGET SOUND REGION. 393

minute ones and gradually increasing in size towards proximal end
of seta.

Nephridiai papillae begin at the 6th somite and extend to 36th ;
with acuminate tips.

Length of largest specimen, 21 mm.; width, including setae,
6 mm.

This species is represented by two imperfect specimens, one from
Puget Sound, collected by Harrington, the other from Coronado,
Cal., collected by me in July, 1895. Its living color is stated in my
notes to be "orange-yellow," but the alcoholic specimen is pale
brown. Elytra and dorsar cirri are easily ^tached. The occur-
rence of the species at localities so widely separated, and, so far
as known, at no intermediate point, is certainly remarkable.

9. Harxnothoe pacifica sp. nov. PI. 2, figs. 14-17.

Form rather flattened, diminishing towards both head and pos-
terior end ; breadth, including setae, about four ninths of length.
Somites, 37; elytra (wanting in specimen), 15 pairs, borne on
somites 2, 4, 5 23, 26, 29, 32.

Prostomiwn very strongly bilobed, nearly twice as broad as
long; "peaks" short and blunt, divaricate; lobes of prostomium
separated by a wide, shallow groove. Basal joint of tentacle very
thick, about the length of the prostomium. Antennae slender,
considerably linger than prostomium, inserted below the level of
tentacle ; ciliate, their basal joints projecting beyond the peaks of
the prostomium. Anterior pair of eyes laterally directed ; just in
front of the bulge of the prostomium ; posterior pair dorsal, near
base of prostomium, slightly nearer together than the anterior.

Peristomial cirri villous, with9ut bulbous enlargement near tip.
Dorsal cirri (Fig. 14) very similar. Dorsal ramus short and
stubby without finger-like process, bearing setae of two different
forms (Figs. 15, 16) which grade into each other. The short
curved ones are the more dorsal. All the dorsal setae are much
more slender than the ventral. Ventral setae (Fig. 17) stout,
hastate, with straight or slightly curved tips, and from two to
twelve serrations. Ventral cirrus fusiform, papillate.

Length^'l^.b ram. ; width across middle, including setae, 11.5 mm.

This species is represented by a single specimen, unfortunately
without elytra and imperfect in other respects. It was collected
by the Columbia Expedition. The species is probably scarce, as all
efforts to obtain more specimens have thus far failed.

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394 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

10. Harmothoe tuta (Grube). PI. 2, figs. 18, 19. PI. 3, figs.
20-22.

Polynoe tuta Grube. Arch. f. naturgesch., jahrg. 21,'bd. 1, 1855,
p. 82.

Haloaydna tuta Baird. Journ. Linn. soc. London, vol. 8 (Zool-
ogy), 1865, p. 188.

Form elongated, of nearly uniform width throughout anterior
two thirds, gradually tapering in posterior third to extremity;
somites numerous and variable in number (77-83) ; elytra not all
paired, 35-38 on each side ; borne on somites 2, 4, 5, 7 ... . 23, 26,
29, 32 .... ; general color of formalin specimens, very pale brown ;
a transverse band of darker brown on every somite of anterior
half, and a median dark stripe.

Proatommm (Fig. 18) distinctly bilobed, median furrow extend-
ing to its base ; lobes rounded anteriorly, widely sundered by the
thick napiform basal joint of tentacle ; eyes placed well forward,
medium-sized, black ; the anterior pair laterally, the posterior pair
upwardly, directed. Basal joints of antennae inserted below the
basal joint of tentacle, on underside of prostomium ; distal segments
of antennae and tentacle very short, nearly equal, not exceeding
length of prostomium. Palpi also short, rapidly tapered to subulate
tips ; constricted at intervals in contraction ; all cephalic appendages,
like tentacular and dorsal cirri, sparsely beset with minute papillae.

Tentacular cirri considerably longer and more slender than ten-
tacle ; subterminal enlargement very slight.

Elytra slightly variable in shape, nearly orbicular (Fig. 20),
some broadly reniform, strongly imbricated, meeting across median
line ; with very few microscopic tubercles, thin and translucent, suf-
fused with smoky brown.

Parapodia (Figs. 18, 19) elongate, nearly or quite equal to
width of dorsum between elytrophores ; dorsal setae rather numer-
ous, much shorter and more slender than the ventral (Fig. 19),
divaricate, slightly curved, rather coarsely serrated along convex
border near tip (Fig. 21). Ventral setae numerous (over twenty),
with about twelve ** frills " near the slightly expanded tip. Point
nearly straight, acute. Dorsal ramus short for this genus, with de-
cided finger-like process, to tip of which the acicula extends. Ven-
tral ramus stout, fleshy, with thick, stumpy, finger-like process which
does not receive tip of acicula. Ventral cirrus short, abruptly taper-
ing to a fine point.

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JOHNSON: POLYCHAETA OF PUGET SOUND REGION. 3^5

Ne^jhridial papiUae extraordinarily long in large specimen, fully
equal to ventral cirri ; begin on the sixth somite and extend to the
eighty-iirst in the larger, and to the seventy-fourth in the smaller
specimen.

Length of larger specimen, 82 mm. ; width, including parapodia
and setae, 12 mm. ; without setae, 10 mm. ; without parapodia, 5
mm. Length of smaller specimen, 61 mm. ; width, including para-
podia and setae, 8 mm. ; without setae, 6 mm. ; without parapodia,
2 mm.

This interesting Polynoid is represented by two specimens in the
collections, one obtained by Harrington and stated by him to be
commensal in the tube of the largest species of ^^Amphitrite " found
by the Expedition, — undoubtedly Thdepus crtspus (see p. 428);
the other collected by Miss Robertson at Alki Point. The latter is
smaller and evidently younger than the former. The dimensions
given in the diagnostic description show dearly the difference in the
proportions of young and old.

The nearly colorless condition, as well as the great length of the
body and the thin, translucent, smooth elytra, indicates plainly its
constantly commensal habit. Forms like P. insignis, P, calif or-
nica,^ and Hamiothoe imbricata^ which are sometimes free-living,
sometimes commensal, retain the pigment, often in heightened inten-
sity, when they have either temporarily or permanently adopted the
latter mode of life.

In spite of the excessive number of somites and elytra, the true
relationship of this form is with the species grouped under the
genus Jfarmothoe (senau extensiori) , and not with Polynoe^ where
numerous somites are much more frequent. The relationship with
Hamnothoe is shown (1) by the structure of the prostomium, (2)
by the finger-like processes of the rami of the parapodia, and (3)
by the sequence of the elytra from the 23d to the 32d somite
(23, 26, 29, 32). A fourth character of less importance is the sparse
papulation of the cirri — a feature almost invariably present in liar-
niothoe^ and absent in Polynoe. The excessive number of somites
and pairs of elytra possessed by not a few commensalistic or parasitic
Polynoids is no doubt correlated with their mode of life ; hence the
unusual length attained by the commensal Ilamiothoe tuta need not
surprise us, although such a length, and elytra in excess of 15 pairs,
seem to be as rare in this genus as they are common in Polynoe,

» SjnoDTm for P. reticulata Johnson which name is preoccupied by P. reticulata
CUparide.

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39«J PROCEEDINGS : BOSTON SOCIETY NATURAL mSTORT.

It is not unlikely that such highly modified fomiB as Poly not
lordly P. pidchra, and P.fragilis have descended from Harnxothot
stock and do not rightly belong in the genos Polynoe, The
characters are a milange of those of Harmothoe and Polynoe (sensu
exL), This may be taken to indicate that these commensals retain
characters of the Polynoid ancestor from which typical Polynoe
and Harmothoe have both descended. The structore of the pro-
stomium, especially the presence of basal joints to the antennae,
recalls Harmothoe \ while the stmcture of the foot is more like
that of Polynoe. The sequence of the elytra differs from that of
both genera. These species certainly do not belong in the genus
Lepidasthenia^ where Darboux ('99) has placed them.

The asymmetrical somites (dorsal cirrus on one side, elytra on
the other) constitute the most striking peculiarity of this species,
and one which, so far as present knowledge goes, it shares only
with Polynoe yigas Johnson ('97, p. 174), JLepidametria com-
mensalis Webster ('79, p. 210), and Polynoe lordi (Baird), In
Polynoe yigas^ I have found at most two unsymmetrical somites;
in P, lordi J from one to nine ; for Lejnd^nnetria the number is not
stated. In two specimens of Hannothoe tuta I have found,
respectively, 15 and 16 asymmetrical somites! As in P. giga^^
they are confined to the posterior part of the body (back of the 32d
somite) whence the sequence of the elytra differs in different
individuals. In the older specimen (Columbia University collec-
tion) there are 38 elytra on the right side and 35 on the left In
the younger 8])ecimen (No. 963) there are 38 elytra on the right
side, 36 on the left. The asymmetrical somites of the former are :
38, 41, 43, 56, 57, 60, 61, 65, 69, 71, 77, 80, 81, 82, 83 — a total of
15 ; those of Xo. 963 are: 33, 37, 3S, 39, 40, 41, 43, 45, 60, 62, 64,
66, 68, 70, 72, 74 — 16 in all.

The identification of the specimens from Puget Sound ^nth the
Pohjnae tift'i of Grube ('55), described from specimens coUected
at Sitka, seems reasonably safe, although his description takes no ac-
count of the diagnostically important structures of the head. The
seciuenoe of the elytra up to the thii-ty-second somite is precisely as
stated by Grube. Beyond that point it differs in every different
individual — a fact not perceived by Grube.

In no Polynoid has a normal asymmetrical somite been found in
front of the 32d.

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JOHNSON: POLYCHAETA OF PUGET SOUND REGION. 397
SiGALlOXIDAE.

11. Sthenelais fusca Johnson.

Sthetielais fusca Johnson. Proc. Cal. acad. sciences, 8d ser.,
Zooloiry, vol. i, 1897, p. 185. Figs. 60, 61, 64.

A single imperfect specimen of huge dimensions, including pro-
stomiuni and 74 somites, was collected by Miss Robertson at Pleasant
Beach in July, 189X. It is far larger than any specimen yet taken
on the California coast. It measures 107 ram. in length (the en-
tire worm must have been over twice as long) and 12 mm. in trans-
verse diameter. The exposed surface of the elytra is rusty, with the
exception of the first six pairs and scattered ones along the body,
which are unpigmented and translucent.

IIesiomdae.

1*2. Podarke pugettensis sp. nov. PI. 3, figs. 23-25.

Body gradually tapering towards both ends ; somites 50-54, the
anterior three or four (Fig. 28) much shortened ; the rest about
four times broader than long.

Pro^toinutiii twice as broad as long, three-lobed in front, the
lobes bearing the tentacle and <lorsal pair of antennae (F'ig. 23) . No
palpi ; antemiae 4, the ventral pair considerably stouter than the
dorsal, and provided with a thick basal segment. Tentacle small,
subulate, less than length of head. E^^es 4, contiguous, anterior
pair slightly largei- and farther apart than the i)osterior; both pairs
with lens ; retinal pigment brown.

Te)itacithtr cirri on first three segments, two pairs to each, with
distinct basal joints; dorsal cirri of 2d and 3d somites equal and
longest.

Panipodia elongated (Fig. 24), exceeding half the width of dor-
sum ; dorsal ramus and basal joint of dorsal cirrus fused ; ventral ra-
mus much longer, terminating in a conical, achaetous tip ; 2 aciculae
in each ramus. Dorsal cirrus notably longer than the parapod and
setae ; gradually tapered ; ventral cirrus extending obliquely back-
ward, somewhat beyond, acute tip of ventral ramus.

ISetae of two sorts, simple and compound, the former confined to
the dorsal ramus; very few (Fig. 24) ventral setae, some with
elongated ap[)endages (Fig. 25) and some with short appendages —
otherwise alike in form. They are arranged in a supra- and an infra _

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39H PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

acicular fascicle, from 13 to 20 setae in each. Anal cirri wanting
in all the specimens.

Leuffth of largest specimen, 25 mm. ; width, 3.8 mm.

A few specimens from Alki Point, collected by Miss Robertson,
Aug. 3, 1898. None are perfect, the tentacular and dorsal cirri, in
particular, being very caducous. The species occurs also at Pacific
Grove.

Xereidae.

13. Nereis virens M. Sars. Pi. 3, figs. 26-30.

Nereis virens Grube. Middendorff's Reise in den aussersten nor-
-den und osten Siberiens, bd. 2, Zool., th. 1, 1851, p. 6. Taf. 1, figs.
2-6.

Alitta brandti Malmgren. Ofversigt af k. vet. akad. f5rhand-
lingar, Stockholm, 1865, p. 183.

N, foliata Baird. Appendix to John Keast Lord's " Naturalist
in Vancouver Island and British Cohmibia," voL 2, 1866, p. 347.

N, brandti Ehlers. Die borsten warmer, 1868, p. 563.

I cannot agree with Ehlers ('68) and Malmgren ('65) that this
North Pacific Nereid is distinct from N, viretis of the coasts of
Northeast America and of Northern Europe. I have compared
specimens from the two oceans, and fail to find any differences that
can be considered of specific value. It is apparently an interesting
instance of discontinuous -distribution, since the species seems not
to attain high northern latitudes. The coast of Norway, the
Gulf of St. Lawrence, and the Sea of Ochotsk (Grube) are the
northern limits of its known distribution. It is likely to be found,
however, on the more southern shores of Alaska. I have not found
it on the California coast further south than Bolinas, about ten miles
north of San Francisco.

The s})ecies attains a much greater size in the Pacific than in the
Atlantic. Whereas an Atlantic xY. virma of a length of 26 cm. in
the contracted condition would be considered large, a length of 50
cm. and a diameter of 25 mm. under the same conditions are not
infrequent dimensions on the western shores of North America
(Fig. 2G).

The Ileteronereis of this species I have not seen, but as usual
among Nereids, individuals attain sexual maturity in the ordinary
or " atokous" form.

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J0HN80N : POLYCHAETA OF PUGET SOUND REGION. 399

14. Nereis vezilloBa Grube. Pis. 8 and 4, figs. 31-88.
Nereis vexillosa Grube. Middendorff's Reise, etc., bd. *i, Zool.,

th. 1, 1851, p. 4. Taf. 2, figs. 1, 5, 6.

iV. arctica Grube. Ibid., p. 11. Taf. 1, fig, 7.

Heteroiiereis midde)idorffii Malmgren. (ifversigt af k. vet. akad.
forhandlingar, Stockholm, 18G5, p. 109.

This species, described and figured a half century ago by Grube,
and subsequently by Ehlers ('68, p. 578), is one of the commonest
Annelids all along the coast, from the Pribylof Islands to Santa Bar-
bara. It apparently abounds in Puget Sound, judging from the
fact that it occurs in all the collections from that region, and numer-
ously in that of the Columbia Expedition. It is known to fisher-
men as the "pile- worm," on account of its habitat amongst the mus-
Bels and barnacles which cluster thickly upon the piles of wharves
and bridges, and is in much requisition for bait. Upon the piles it
seldom attains a length greater than 20 cm. and a diameter of 12 mm.;
but in gravelly beaches, where it lives in company with iVI virens,
it grows to a somewhat larger size. Olive-green is its usual color
in life, but this changes to a bright emerald-green or bluish green in
alcohol. Tints of brown are very frequent in the more posterior
parts, and sometimes the entire worm is brown or dusky.

The great dorsal lobe of the feet of the posterior portion of the
body, carrying at its distal end the dorsal cirrus, increases notably
in length with age, as may be seen by comparing Figs. 84 and 85,
PI. 4. The former represents a foot of a young female; the
latter, the foot of a large, sexually-mature female. This great
increase in length is probably not correlated with the advent of
sexual maturity, as some individuals of very diminutive size have
the dorsal lobes of considerable length. In passing caudad the
dorsal lobes lengthen so gradually it is impossible to find a demarca-
tion-point between long and short lobes. In the anterior region
in front of the 12th foot the feet are of the form shown in Fig. 81,
PL 8 (tenth foot).

The Heteronereis (epitokous form of Ehlers) is not uncom-
mon (Figs. 32, 83). This condition is by no means confined to full-
grown worms, but occurs in individuals having a length of 56 mm.
and upward. The species also arrives at sexual maturity without
becoming heteronereized, as I have noted in many instances.

15. Nereis agassixi Ehlers. PI. 4, figs. 89-45.

Nereis agassizi Ehlers. Die borstenwttrmer, 1868, p. 542.
PI. 28, fig. 1.

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400 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

This beautiful Nereid is a tube-dweller. The secretion is fur-
nished by the large glands opening on the dorsal aspect of the
upi)er lobes of the feet (Figs. 44, 4.iy, PI. 4) , and is poured out so
rapidly and cof)iously that a new tube is formed in a very few
minutes when a worm is removed from its tube and placed in a jar
of clean sea-water. The tube is rather tough and leathery, but
very flexible and translucent, especially when newly secreted. The
worms are gregarious, and great tangled ropy bunches of ** eel-
grass'* {Plnjllipsjf*idi,r) are often found, held together by the power-
fully-adhesive secretion of a colony of X. mjtissizi. The species
attains sexual maturity at the latitude of San Francisco in February
and March. I have rarely seen the Ileteronereis of this species.

X. iitjKH.'iizi docs not attain a hirtre size. The largest I have col-
lected measures 08 mm. in length and .') mm. in diameter, including
parapodia and setae. Several small specimens have been brought
from Puget Sound, and I have collected it at various points along
the California coast as far south as Santa Barbara.

10. Nereis cyclnms Harrington. PI. 4, fig. 46. PL 5.
figs. 48-52.

X* rets ci/chirus Harrington. Trans. X. Y. acad. sciences, vol.
10, IS97, p. :>14. Pis. 10-18.

"^riiis remarkable species has been ably described by Harrington
('97), and its extraordinary commensalistic relations with the Her-
mit crab { Eiip'KjUrns iinnntas Dana) are discussed.' The scanty
material at command <loes not enable me to add an^-thing of value.
One of the striking features of this Nereid is the immense size and
cup-like form of the peristomium (Figs. 40-48). This is undoubtedly
for the protection of the prostomium, which when retracted is par-
tially concealed within the concavity of the peristomium.

17. Nereis procera Ehlers. PI. 4, fig. 47. PI. 5, figs. 58-59.

Xcrels i>r()Ctt'ti Ehlers. Die borstenwtlrmer, 1808, p. 557.
PI. -28, tig. 2.

A very slender Xereid obtained by the Columbia Expedition
un<loul)tedly belongs to this species. There are several specimens,
none of which is <'om])lete. The species does not occur in the
other collections from the Sound. There are no data as to depth
or locality.

It is highly probable that this is the species of Xereis mentioned
by Harrington and Griflin ('97a, p. 150) as dwelUng in the tul>es of
a Chaetopterid. The extremely attenuated form of the body and

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JOHNSON: POLYCHAETA OF PUGET SOUND REGION. 401

great number of somites suggest such a habitat ; and the stout
dorsal seta shown in PI. 5, Fig. 59, is such as would properly belbng
to a tube-dwelling Nereid. These setae remind one of the " hooded
crotchets" of Nereis agassizi (PL 4, Ftg. 43), which is also a tube-
dweller, but they are in fact compound setae with a much reduced
appendage deeply sunk within the tip of the shaft. These setae
are not found in the most anterior parapodia, but begin about the
40th foot. A seta of this form would be especially useful in clam-
bering within the tube. These stout setae are only one or two in
number in each foot (Figs. 54, 55, 56), and occur only in the dorsal
ramus. There are no setae of the ordinary form in the dorsal
ramus where these setae are present, except in two or three
parapodia where the change is taking place (Fig. 54).

Ehlers's description of this species is based upon a single specimen
collected by Alexander Agassiz in the Gulf of Georgia in 1859.
This type, now deposited in the Museum of Comparative Zoology,
I have had an opportunity to ejcamine, and I find it of the same
species as the specimens above mentioned.

None of the specimens is complete. Ehlers's example had a
length of 125 mm. and 179 somites; a nearly perfect specimen in
the Columbian collection measures 146 mm. and has *260 somites.
It is evidently not full grown, for its greatest transverse diameter,
including parapodia, does not exceed 3 mm., whereas Ehlers's speci-
men, which is the largest I have seen, has a diameter of 4 mm.

Nephthydidae.

18. Nephthys coeca (Fabricius) ()rsted.

Numerous examples from various localities — Neah Bay, Salmon
Bay, and Pleasant Beach. The largest specimens measure *20 cm.
and over in length, and 15 mm. across the thickest portion, including
the parapodia. The 8i)ecies occurs northward along the Alaskan
shores, in Bering Sea (Marenzeller, '90) and along the northern
coast of Siberia (VViren, '83). It extends southward along the
California, coast as far at least as San Francisco Bay, but the Cali-
fornia specimens are pygmies as compared with those from Puget
Sound and Alaska.

The species was long ago collected by Alex. Agassiz in the Gulf
of Georgia and identified by Ehlers (" Die borstenwttrmer,'* p. 588)
as identical with Nphthys coeca of European waters. I have com

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402 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

jiared the Puget Sound specimens with sopie in the Museum of
Comparative Zoologj' from Massachusetts Bay, and find them iden-
tical in every respect.

EUPHROSYNIDAE.

19. EuphroBjrne heterobranchia sp. nov. PI. 6, figs. 60-66
</-c.

Form elliptical, robust, slightly more tapering towards the pos-
terior than towards the anterior extremity. Dorsal bare stripe
narrow, less than one fourth the width of body. Somites, 34.

Carnnde (Fig. 60) low, bilobed dorso-ventrally, the lobes of
ecjual length, reaching sixth somite. It has eight longitudinal
ridges, two pairs in the upi)er, and two in the lower lobe, extending
the entire length of the caruncle. Median tentacle short and awl-
shape<l, its filiform tip nearly as long as the thick basal portion ; at
its base the two posterior or '* dorsal " eye-spots. The anterior or
** ventral " eye-spots confluent, flanked on each side by a very minute
antenna.

palpi rather broad and flat, separated by a slight furrow from first
and second somites, divided by a conspicuous median cleft. The
mouth bordered posteriorly by the 5th somite.

Punipiulin of usual form in this genus. A short ventral cirrus at
posterior edge, adjacent to the intersegmental furrow ; a lateral cir-
rus between the fourth and fifth gill-trunks, counting from the upper-
most of the series ; a stout, fusiform doreal cirrus, not exceeding the
branchiae. Branchiae ten on each side, some simply forked, others
ramose, branching twice (Figs. 66 a-c). Setae all with hard, glisten-
ing tips ; the bitid ones of doi*sal series often have a very minute
lateral tooth (Fig. 63); lateral tooth of ventral series also small
(Fig. 65); cleft setae both serrate and non-serrate (Figs. 61-62<0,
the latter form the more common ; serrations sometimes very few.

Jjfn^jth^ 18 mm.; width, not including setae, 4.5 mm.; dorso-
ventral thickness, 88 mm. ; median base stripe, 1 mm.

A single specimen in the Columbia collection. The species is
interesting for its resemblance to E. horedlis of the North .Atlantic,
as to the heterogeneous character and large number of its bran-
chiae, but differs from it in the shape and multiform nature of
its setae, and in the larger number of somites. It is sufficiently dis-
tinct from all other known species of our Pacific coast, although its
superficial resemblance to Enphrosyne urctia is rather striking.

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JOHNSON: POLYCHAETA OF PUGET SOUND REGION. 403

As regards the so-called ** branchiae " of Euphrosyne^ it was long
ago pointed out by Clapar^de ('68, p. 420) that functionally these
are no branchiae, but probably glands, which furnish the slime with
which the animal is more or less coated. I have recently had an
opportunity to examine these structures in a young specimen of
E, atmtntidca and compare them with the functional branchiae of
Etwythoe californica^ both in situ and freshly abcised from the liv-
ing animal. While the branchiae of Eurythoe are richly vascular,
the organs of Euphrasy ne show not the slightest trace of blood-
vessels, although almost as transparent as those of Eurythoe. On
the other hand, these gill-like structures are richly beset with gland-
ular cells, and there cannot be much doubt that they furnish a part
or the whole of the mucus with which the setae are usually enslimed.
The relationship of Euphrosyne to Eurythoe is undoubtedly close ;
many authorities place them in the same family. We may therefore
legitimately conclude that here we have homologous structures that
have undergone a change of function; and the term "branchia,"
applied in a morphologic sense, is not a misnomer, although perhaps
liable to be misleading.

Study of sections of specimens of Euphrosyne (lurnntioco and
Eurythoe c<ilifor7iic<(^ fixed in aceto-sublimate and stained with
haemalum, shows less difference in the structure of the gills than
would appear to exist from examination of the structures in the liv-
ing or fresh condition. Both, indeed, contain blood-vessels or struc-
tures functioning as such. In Eurythoe the presence of a vascular
loop is very evident, but in Euphrosyne I have not been able to
detect a loop, or in fact anything more than a cleft in the tissue
sometimes eni]>ty, sometimes tilled with coagulum. This seems to
be a lymph space connected with the body cavity. In both species
the walls of the gills are thick, but thicker in Euphrosyne than in
Eurythot', The branchiae of Euphrosyne contain an axial strand of
muscle fibres. Their surface is ciliated.

SVLLTDAE.

20. Pionosyllis elongata sp. nov. PI. 6, fii^s. 07-70. PI. 7,
fig. 71.

/^or/?i slender, becoming much elongated with age; 140-200 so-
mites ; diameter nearly uniform the entire length, tapered slightly
towards head and tail ; intersegmental furrows are deeply incised ;
somites average two^and one half times as broad as long.

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404 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

Prostomiam (Fig. 67) about one and one half times as broad as
long, in front obtuse, convex, the base nearly full width of peristo-
mium. Palpi fused for nearly half their length. Antennae and
median cirrus moniliform, similar to peristomial and dorsal cirri.
Eyes four, the anterior pair twice the size of the posterior, and con-
siderably further apart.

Periston! i am with dorsal and ventral cirri, without parapodia or
setae.

Par<ipo(lia (Figs. 67, 09) uniramous, with double aciculae; setae
few; ventral cirri short, plain; dorsal cirii moniliform, gradually
tapered from the l)a8e; slightly exceedhig in length the diameter of
the worm. Articulations 10-15 in antennae and peristomial cirri,
increase to 16-18 (in some specimens, 18-20) in dorsal cirri of the
antenor region, then diminish to 14-15 in more posterior somites.

Pyifidhjin (Fig. OS) hemispherical ; anus on its dorsal side ; anal
cirri longer than the dorsal cirri, cylindrical, 16-17 jointed.

AUiiteritary atnal (Fig. 71) of usual form ; proboscis with a
circlet of thirteen conical papillae (pop,) at its orifice, and armed
with a single tooth (f.) near anterior edge of chitinized lining; pro-
boscis frequently found everted in preserved specimens. Oesopha-
gus (oes.) elongated, extending through about twelve somites ; its
posterior third thick-walled and glandular. The proventriculus or
"gizzard" (prov.) of the usual structure, extending through eight
to ten somites, according to degree of contraction of body ; cyhn-
drioal, rounded at the ends, lumen verj' narrow. Ventricular coeca
(c.) much elongated, extending through four somites, frequently
curved or bent double and opening into digestive tube just back of
the proventriculus.

>SVa*fc.v distinct; no stolonization ; genital products develop only
in posterior somites (10))d to lO-ltli in a female specimen with 198
somites), wliicli become much enlarged in consequence.

(/oh)/\ in life, nearly white, translucent; ova rich yellow.

Leniith of full-grown female (19>< somites) 5S.5 mm.; transverse
diameter, iucludinii: parapodia, 1.1 mm.

A single sjiecimcu was collected by Miss Rol>ert8on at Port
Orchard in July, ls99. It is immature, measures 31 mm. in length,
and has but 140 somites.

This species occurs within tide-marks as far south as Pacific
Grove, Cal, where I have taken sexually mature specimens in De-
cember. I have also collected, in the month of February, sexually

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JOHNSON: POLYCHAETA OF PUGET SOUND REGION. 405

mature specimenB of exceptionally large size at Point Cavallo, on
the northern shore of the Golden Gate, where it occurs in small
numbers on the underside of stones, inside of dead shells, etc. The
much-swollen caudal segments, turgid with ripe ova, are very con-
spicuous.

I have examined the radial muscle-columns in the proventriculus,
and find the structure as regards the central core of granular, undif-
ferentiated protoplasm, in which the nuclei are located, and as
regards the peripheral layer of muscle tissues, in harmony with the
statements of Haswell (*86) and Malaquin ('93); I have not, how-
ever, been able to discern striations in the muscular part, but this
may be due to the fact that the material was not preserved with a
view to histological study.

21. Trypanosyllis gemmipara sp. no v. PI. 7, figs, 72-76.

Form elongated, much flattened dorso-ventrally, tapered towards
both ends, abruptly towards the head, gradually towards the
pygidium ; somites very short ; parapodia less than one sixth the
width of the trunk in its widest portion ; somites very numerous
(300 or more).

Prostotniurtt (Fig. 72) comparatively small, broadest in front,
distinctly bilobed, the lobes separated by a median fun*ow ; eyes
four, the anterior pair larger and very slightly further apart than
the posterior. Median cirrus nearly twice as long as the antennae ;
these, as also the peristomial cirri and all the dorsal ciiTi, monili-
form, with numerous short articulations, diminishing in size towards
the tip. Entire surface of cirri covered with dark brown, easily
detached, bud-like bodies (Figs. 72 and 74). Palpi (^>.) reniform,
elongated, projecting far in front of the prostomium, widely sepa-
rated their whole length.

Peristominui extremely short, embracing the prostomium on its
two sides ; bearing at its anterior corners two pairs of forwardly
directed peristomial cim, of which the dorsals are twice the length
of the ventrals. The arrangement of prostomium and peristomium,
together with their appendages, closely resembles the collocation of
these parts in the Polynoids.

Parapoilia (Fig. 74) not prominent, ventral ramus fairly devel-
oped, pointed at tip, with 7-9 setae of the form shown in Fig. 75 ;
a small separate lobe covers the tips of the double aciculae. Ven-
tral cirrus (v. c.) short, often curved, blunt at tip, non-moniliform.
Dorsal cirrus very long, either straight or circinate at tip ; in the

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406 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

latter case incurved over the dorsum, in the former extending
nearly at right angles to longitudinal axis of the body.

iJiijestive systtjn exhibits a well-developed trepan (Fig. 73) of
ten teeth ; these are surrounded by a circlet of elongated papillae
(pep.). Oesophagus of moderate length (extending through 22
somites), strongly chitinized ; proventriculus cylindrical, of uniform
diameter throughout; two well-defined lateral raphes ; radial muscle-
columns very large. Alimentary canal back of proventriculus with
extensive, segmental, paired diverticula.

Dor Aid surf are elegantly marked with ^ne transverse dark lines
which indicate the boundaries of the segments.

Posterior extroiitty cajiable of producing successive crops of col-
lateral, sexual buds or zooids (Fig, 7(5), which possess every exter-
nal structure of the parent except mouth and anus. They lack,
however, an alimentary canal and nephridia.

Le)ujthy BS mm. ; transverse diameter, 3 ram. ; dorso-ventral
diameter, 1 mm.

A single specimen was collected by the Columbia University
expedition, probably in the vicinity of Port Townsend. Unfor-
tunately, no data accompany it. This individual possesses the
remarkable sexual zooids, over fifty in number, presenting all
stages of development. They arise as collateral buds from a prolif-
erating somite near the posterior extremity (Fig. 76). At full
maturity they evidently separate from the asexual stock and become
frec-swimminix sexual zooids, provided with para])odia, antennae,
eyes, and central nervous system, but destitute of an alimentary
canal. Tliey will be fully described in volume 2 of the Biological
Bulletin.

OXI PHIDIDAE.

22. Northia elegans sp. nov. PI. s, figs. 77-85.

Form stout, flattened dorso-ventrally, tapering towards head ;
except in most anterior region, dorsal contour flattened and ventral
convex ; branchiae simple, filiform, upraised, and slightly incurved
over the back.

ProstoiiuKm (Fig. 77) small, conical, considerably broader than
long, and its sui-face monopolized by its large appendages ; three
pairs of antennae ; most anterior j)air short, ovoid ; second pair with
annulate basal joints and acute tenninal joints about one half the

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JOHNSON: POLYCHAETA OF PUGET SOUND REGION. 407

length of the basal joints ; third pair three-jointed, the middle joint
much the shortest, and the distal twice as long as the proximal ;
this pair more than twice as long as the second pair, and reaching
the sixth somite. Basal joints of second and third pairs of antennae
with nine or ten obscure annulations. Median cirrus likewise three-
jointed, one half as long as third pair of antennae, and with long
terminal stylode ; its basal joint five-annulate. Palpi large, globose,
approximate (Fig. 78,/>.). Eyes four, small, at bases of third pair of
antennae, one pair directed forward, the other laterally (Fig. 78).

Peristomium (Fig. 77, 78) shorter and narrower than the other
somites, having on its antero-dorsal border a pair of small, slender,
peristoraial cirri (Fig. 78,/>. c).

Somites : first four or five back of peristomium longer and nar-
rower than the succeeding ones, with parapodia (Figs. 77, 80) of
different form from the rest, characterized by an elongated ventral
ramus, with acute achaetous terminal portion, a large fusiform
ventral cirrus (y. c), a dorsal cirrus {cL c.) of similar form and
dimensions, and a cirriform gill {br.), likewise of similar aspect.
The transition to the typical somite and foot is gradual {cf. Fig.
81, 28th foot).

Branchiae unbranched throughout, tapered to an acute tip,
increasing in length caudad, until they exceed half the transverse
diameter of the trunk. The setae of the 4-5 anterior parapodia are
different from those of the succeeding feet, and are of two forms,
"hooded crotchets" and capillary bristles (Figs. 82, 83). Setae
of dorsal rami throughout the series are buried in the foot ; at
most, their tips protrude (Fig. S5) . Setae of ventral rami beyond
fifth foot are (1) bordered capillary (Fig. 84), (2) a pair of stout,
two-pronged uncini with flabellar expansions at tip — in all respects
like uncini of succeeding species (Fig. 90) .

Upper jaws (Fig. 79) asymmetrical, six pieces on right, seven on
the left.

LeiKjth of 85 somites, 66 mm. ; transverse diameter, 6 mm.

The antero-dorsal portion of the trunk eleijantly marked with
paired umber-brown spots })laced near the posterior border of each
somite ; these tend to coalesce across the median line (Fig. 77) .

Three or four specimens of this fine Northia occur in the Colum-
bia University collection. Unfortunately, all lack the posterior
portion of the trunk. There are no data as to depth or exact local-
ity. The tubes also are lacking.

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408 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

23. Northia irldescens sp. nov. PI. 8, figs. 86, 87. PI. 9,

figs. 88-92.

Form slender and nearly cylindrical, of almost uniform diameter
in anterior region ; highly iridescent ; branchiae curved over dor-
sum, not reaching median line ; slender, filiform, translucent (Fig.
88).

ProHtoitiium (Figs. i^G, ><S) small, hemispherical; first or "acces-
sory " antennae elongate ovate, constricted at base ; second pair of
antennae two-jointed, basal joint slightly the longer, twelve-ringed ;
third pair of antennae three-jointed, with 11-1-8 annulations; this
pair nearly four times as long as the second pair. Median cirrus a
little shorter than third pair of antennae; nine-ringed. Eyes want-
ing. Palpi (Fig. 80) globose, pedunculate, spreading.

Peristfunitutt much broader than prostomium, with a pair of
minute peristomial cirri (Fig. x>^) projecting from its anterior bor-
der.

Jaics (Fig. 87) very similar to those of K. degans^ but with
more teeth on the dental plates and more slender maxillae.

Somites: four following the peristomium with modified parapodia
(Fig. 88), similar to the same somites in X, elegatis; the succeed-
ing ones with filiform gills and dorsal cirri, but no ventral cirri (Fig.
89).

Hooded crotchvtfi (Fig. 91) of ventral fascicles of first four
somites very similar to those of preceding species. Two stout, wing-
tipped uncini (Fig. 90) in ventral fascicles of parapodia further
back. Capillary setae with striated border (Fig. 92) .

Lt^ngth of f)2 somites, 38 mm. ; transverse diameter, 3 mm.

This species is represented by a single specimen, dredged b}" Prof.
W. .V. Herd man at Victoria, B. C, in the summer of 1897, and by
him kindly placed in niy hands for description. Unfortunately, all
the posterior portion of the specimen is lacking. The tube is of
parchment-like mateiial, opaque-white, flexible, and with adherent
sand-grains.

LUMBRICOXEREIDAE.

24. Lumbriconereis zonata sp. nov. PL 9, figs. 93-100.

Form cylindrical, slightly tapered towards anterior end ; para-
podia placed a little below the mid-lateral line ; somites three times
as broad as long, each marked with a sharply-defined brown zone

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which extends around the trunk, widening on each side, above and
below, as it approaches the parapodiuni. Middle of each band
marked by a sharp dark line (Figs. 93, 95, 96).

Prostotaium (Fig. 93) acorn-shaped, of a lighter tint than the
trunk ; perfectly smooth and glistening.

Peristoiniuni (Fig. 93) tapered to the diameter of the base of the
prostomium which is less than three fourths the diameter of the
third somite. Second somite about one half the average length of
the somites.

Jtttrs as shown in Fig. 94.

Farapodia (Figs. 97, 98) less than one half diameter of body,
bi-lobed at tip, posterior lobe the longer. Setae inserted between
the lobes. Setae, as usual, of two forms : winged capillary (Fig.
99) in antenor portion of body and ''hooded crochets" (Fig. 100)
in the posterior portion.

Length of *200 somites, 1()7 mm. ; diameter, including parapodia,
4.5 mm. ; without parai)0(lia, 3 mm.

A single imperfect specimen lacking the posterior region, was col-
lected by Professor Ritter at Salmon Bay, Puget Sound, May 29,
1><99. This specimen is remarkable for the possession of abnormal
segmentation in as many as five places. In two instances the so-
mites are spiral in the way shown in Figs. 95, 9i), representing re-
spectively the dorsal and ventral sides. In another place, the spiral
extends through nine somites, with a forked somite at each extremity.
The other two instances are partially-divided somites without a spiral
arrangement — in one case with the parapodium displaced towards
the ventral side. The asymmetrical somites are not confined to
any limited region, but are scattered for a long distance through the
middle region of the body.

Glyceridae.

25. Olycera rugosa sp. nov. PI. 10, figs. 101, 102.

Form stout, terete, thickest about one third the distance from
head to posterior extremity, tapering slightly cei)halad ; much more,
though gradually, caudad. Number of somites 200-300, distinctly
two-ringed, all setigerous except the pygidium. The rings are
nearly equal, but the anterior one, which bears the parapodia, is often
raised like a welt, giving the body a corrugated aspect.

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Prostomium conical, tapering, 2-3 mm. long, length exceeding
its breadth, obscurely twelve-ringed ; four minute tentacles at tip.
Basal portion not sharply set off from the peristomium. Palpi (?)
retracted.

Pygidium small, globose, bearing a pair of slender subulate anal
cirri. Anus minute, on dorsal side of pygidium.

Parapodia (Fig. 101) stout, of nearly equal height and length,
the largest equal to width of dorsum in its widest part ; except the
two most anterior pairs, four-lobed, the two anterior lobes slightly
longer than the posterior, but all lobes of nearly same form
except in most anterior parapodia. Anterior dorsal lobe sometimes
bifid. All the lobes more or less conical and pointed y their tips
darkly pigmented ; simple capillary setae inserted between the dor.
sal lobes ; the compound setae between the ventral. Ventral cirrus
{v, c.) large, conical, strongly resembling the lobes; dorsal cirrus
{d, c.) smaller, globose, much constricted at the point of attaclmient,
placed high, at the base of the parapod.

Branchiae (iP'ig. 101) begin at the 16th or 17th parapod and ex-
tend to the 22d from the pygidium ; they consist of eight or nine
finger-like, thin-walled lobes, sometimes bifurcated ; completely re-
tractible into body-wall ; at sexual maturity crowded with reproduc-
tive cells (Fig. 101). All branchiae are on posterior aspect of the
parapodium ; the most anterior and most posterior of the series are
single, sausage-shaped processes.

Proboscis extremely variable in length (12 to 35 mm. or more),
club-shaped, thicker than anterior portion of body, beset with mi-
nute papillae of two forms, conical and ovate. Jaws (Fig. 102)
strongly hooked; each bears a triangular appendage (<//>.) with
long falcate process.

Color of alcoholic and formalin specimens variable, from tawny
or buff to olive-brown. The color is due to abundant yellowish
brown pigment grains in tlie liypoderrais. These are often ag-
gre.ijated towards tips of the lobes of the parapodia.

Lenyth of lartre specimen (much contracted), 170 mm. ; transverse
diameter, inchiding parapodia, 9 mm.

This species is probably abundant in the Puget Sound region,
as it occurs in all the collections. It has been taken at Neah Bay,
and at Salmon Bay (near Seattle) ; and there are a considerable
number in the Columbia collection ( probably from the vicinity of
Port Townsend) for which no locality is given. A Glyoerid col-

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lected by J. K. Lord at Esquimalt and described under the name
Glycera corrugata by Baird ('63) is probably identical with G.
rugosa, but the description is too meager to admit of positive deter-
mination. Many of the larger specimens are females with ripe or
nearly ripe ova ; but I have seen no specimen which has undergone
the atrophy of the proboscis incidental to sexual maturity mentioned
by Arwidsson ('98, p. 6).

As the Glyceridae are destitute of a vascular system, the gills are
merely reversible pouches of the body- wall, into which the caelo-
mic fluid passes. Reproductive cells also enter these thin-walled
pouches.

26. G-lycera nana sp. nov. PI. 10, flgs. 103, 108a.

Form short, thick, somites comparatively few (about 140 in one
specimen) ; diameter nearly uniform for greater portion of length ;
somites two-ringed ; the posterior ring slightly raised above the level.

Pro8t(yniium conical, ten-ringed, four minute tentacles at tip. Pro-
boscis short, club-shaped, beset with conical papillae. Jaw-appen-
dage as shown in Fig. 1 08a.

Parapodia (Fig. 103) rather slender, anterior lobes two, the
ventral one the longer, both conical ; posterior lobe single, rounded ;
ventral cirrus (w. c.) similar in shape to upper anterior lobe ; dorsal
cirrus a rounded tubercle placed high above the foot on the side of
body ; no gills.

Setae elongate, of the usual two forms, capillary dorsal, and com-
pound ventral.

Length of larger specimen, 64 mm. ; greatest transverse diameter,
6 mm. ; without parapodia, 4 mm.

The species is present in the Columbia collection and also in
Miss Robertson's; she obtained it at Port Orchard in July, 1898.
The exact locality is not recorded for the other specimen. In both
examples the posterior portion is regenerating, so it is impossible
to give accurately the normal number of somites. It probably lies
between 180 and 200.

27. Hemipodia borealis s[). nov. PI. 10, figs. 104, 104a.
Form terete, moderately long and slender, of nearly uniform

thickness for the greater portion of the length, but tapered poste-
riorly; somites three-ringed, 126 in number; two minute anal cirri.
Prostomunn with conical, ringed process, tipped with four (?) ten-
tacles. Proboscis beset with minute, ovate papillae. Jaws with
notch near base; jaw-appendage (Fig. 104a) a simple rod.

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412 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

Parapmlvt (F'ig. 104), as invariably in this genus, with only the
lower ramus, a single acicula, and no simple setae. The dorsal
cirrus {(J. r.) near the foot, ovate ; no gills. Parapodia consisting of
an anterior, elongated lobe and a posterior, short, rounded one.

LiHf/th, I'l mm. ; transverse diameter, including parapodia, 8 mm.

Only a single female specimen of this ver\' interesting sj)ecies oc-
curs in the Pnget Sound collections. It was gathered by the Co-
lumbia I'niversity Exj»e<lition — exact locality not known. It con-
tains large cltl'^s, and is evi<lently mature.

The gtnus IleiHiinul'm was established in 1SG5 by <Juatrefages
for the reception of a ])eculiar Glycerid from Chile, which he named
11. ro.st//.v. The same year another species fi*om the StraitiS of
Mairellau was described uniler the name //. patafjontaf by Kinberg
('Go, p. !i4.">). I'ntil tiie ])rese]it, no genuine species of IhinljuHlin
has been added to the two original (uies, although two sj>ecies of
Glyceridae — y/e//< />''><///' (?) m(t>/*Itanir(i M'Intosh ('So, p. 349)
and //. st-j'f^'ttfi'lnmifis Roule ('IHI, j). 4')*2) — have been erronef»usly
attributed to tliis genus {ritJe Arwidsson 'OS, p. 28) .

.\UH IIDAK.

*is. Scoloplos elongata sp. nov. PI. 10, figs. 105-110.

Form lonir and slender, somites short and very numerous ( 298 in
one s])ecimen) ; tlattcne<l anteriorly ; broadest between 9th and
17th somites ; thence narrowing gradually to a uniform diameter
which is kept about as far as the 200th somite, thence gradually and
unifonnly diminishing to the slightly expanded, hemispherical py-
gidium. Dorsum plane, but a])parently concave on account of the
upward direction of the i)arapodia. Ventral aspect convex, flattest
in the widened anterior region, where dorsal and ventral surfaces
are nearly alike and the parapodia are laterally directed.

Pr<>i<fn,t,hi)n small, tip)»ed by a conical, acutely-pointed palpode
(Fiic. 1 <>.'), pp.) ; without eyes.

Peristuiitinin increasing rapidly in width towards the second
somite, which is the first to bear setae. Pharynx eversible, with
leaf-shape<l lobes (Fig. 100).

Purdj-iotlin at anterior end (Fig. 107) with dorsal and ventral
setae, an<l sim])le branchiae. Setae closely serrated CFig. 109).
Parapodia back of the anterior region (Fig. 108) with larger, ciliated

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JOHNSON: POLYCHAETA OF PUGET SOUND REGION. 413

branchiae and dorsal cirri. Dorsal and ventral setae in small fas-
cicles. Ventral ramus of foot continued ventrad as a leaf-like ex-
pansion, but not extending below the medio-lateral line. Anus
on dorsal side of pygidium ; anal cirri long and slender.

Lengthy 192 mm. ; width of thorax, 3 mm. ; width of abdomen
(including parapodia), 2 mm.

This species burrows in sandy shores, near low- water mark,
whereas the much commoner and more widely distributed species of
Aricia found on the Calif ornian coast has its habitat among the
rhizomes of Phyllospadix, The present species is represented in
the material from Puget Sound by a number of incomplete speci-
mens collected by the Columbia University Expedition and by Miss
Robertson.

Levinsenhdae.

Aricideopsis gen. nov.

Characters of Ariciden Benedict and Webster, but differs from
it in having well-developed parapodia and setae on the peristomium,
and no ventral cirri. Eyes very large, compound. Anal cirri, two.

29. Aricideopsis megalops sp. nov. PI. 10, figs. Ill, 112.
PI. 11, figs. 113,114.

Form of moderate length, gradually tapering from the anterior
to the posterior extremity, color of preserved specimen pale yellow,
the dark intestine showing through in the posterior portion. Para-
podia with distinct rami flattened into foliaceous expansions, much
the largest dorsally, and in the anterior half of the body ; setae in
both rami; uncini and capillary setae in all ventral rami (except
the last two or three, which are achaetous) back of the 18th.

Proatomium (Fig. Ill), the full breadth of trunk, rounded ante-
riorly, elevated in a median thickening at base, which is bilobed
anteriorly, and bears the median cirrus and the large crescentic
eyes.

Parapodia (Figs. 112, 113) with dorsal rami expanded into leaf-
like form on somites 2-24 and V>eyond, gradually becoming smaller
and more rounded ; towards end of series the upper lobe becomes
minute ; ventral lobe rounded, always smaller than the dorsal ; a
filiform gill {br.) on somites 2-27, often with lanceolate tip. Setae
(Figs. 112, 113, 114) of two sorts : capillary and " hooded crotchets " ;
both kinds occur in the ventral rami back of the 17th somite; at

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414 iraOCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

f&nrt'Only one or two uncini (Fig. 114) among the capillarj bristles,
;gra(}ua1!ly increasing to ^five, while the capillary bristles decrease
pari passu. Anal cirri two, short and stumpy.

The present species undoubtedly belongs to the family LeTin-
seniidae recently established by Mesnil and Caullery ('98) for the
reception of a small group of peculiar little Polychaetes which show
affinities to the Spionidae on the one hand, and to the Ariciidae on
the other. The species under consideration most neafly resembles
Aricidea, of which two species have been described from the Atlan-
:tic coast by Webster and Benedict ('87). It differs enough how-
ever from Aricideft to deserve generic distinction. The presence of
parapodia and setae on tlie peristomium probably indicates a more
primitive character than the allied genus exhibits. The large size
of the eyes is also remarkable, and would seem to indicate a pelagic
!habit.

The foregoing description is based upon a single specimen from
Port .(Orchard, jL'ollected by Miss Robertson in June, 1899.

Magelonidae.

30. MagfilDna longicomis sp. nov. PI. 11, figs. 115-118.

Fomn cylindrical, rather stout, of nearly uniform diameter,
divided iatio two regions: (1) the ajiterioTy in front of ninth
somite, with capillary, double-bordered setae (Figs. 116-117) in
both fascicles ; (2) the posterior, back of and including the ninth
somite, with uncini (Fig. 118) both above and below. Ninth
somite (Fig. IIG) shorter than the others, and marked by a
deeper constriction, with a pair of comb-like fascicles of short, stiff,
iiapillary setae. ^

Prostornimn (Fig. 115) flattened, grooved in median dorsal line,
anterior tip exj)anded ; no eyes. Peristomium beat's a pair of
enormously long, flexile, tentacular cirri, beset with numerous cap-
itate j)apillae on exterior aspect; showing rings of contraction near
its base (Fig. 115). Proboscis (/>r.) rounded, without corrugations
or surface differentiation ; extensible as far as tip of prostomium.

Purnpodia (Figs. 115, 116) of anterior region slightly developed;
dorsal and ventral cirri small; a small branchia between them;
dorsal and ventral setae (Fig. 117) of same form, double-bordered
capillary. In posterior region, branchiae and cirri are larger ;
uncini (Fig. 118) inform of "hooded crotchets" bidentate at tip,
in transveree rows of ten or eleven, rising high upon dorsal side.

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JOHNSON: POLYCHAETA OF PI GET SOUND REGION. 415

Length of head and anterior region, 7 mm. ; length of first 12
somites of abdomen, 8 mm. ; greatest transverse diameter, 1.5 mm.;
length of tentacular cirri, about 14 mm.

Two imperfect specimens, consisting of only a few anterior
somites, represent this interesting species. It was collected by
Miss Robertson at West Seattle, June 23, 1899.

Magdona papiUicoruia^ originally described by Fritz MttUer ('58)
from the Island of Santa Catharina off the coast of Brazil, has
since been found on both sides of the North Atlantic (vide
Andrews, '91). Its anatomy, both external and internal, has been
carefully studied by M'Intosh (78) and its remarkable blood has
been investigated by Benham ('96). Hitherto it has remained a
unique and isolated form, most closely related to the Spionidae but,
as M'Intosh pointed out, having affinities also with the Chaetop-
teridae. The present species differs from J/ pfipillicomis (1) in
it« much greater size, (2) in the gi'eater length of its tentacular
cirri and longer papillate areas of same, (3) in the comparative
shortness of the prostomium, and (4) in the smoothness of the
proboscis.

Capitellidae.

31. Capitella dizonata sp. nov. PI. 11, figs. 119-121.

Thorax thickest in region of 5th and Gth somites; smallest at
8th and 9th (Fig. 119), most of the thoracic somites two-ringed;
abdominal somites three- to twelve-ringed; intersegmental con-
strictions pronounced, especially in thorax.

Prostomium short, conical, at base slightly more than one half
the diameter of the peristomium; nuchal organs not discovered.

Peristomium setigerous ; somites of thorax over three times as
broad as long in the contracted state ; the 4th and 5tli each with
a dark brown band passing around it in front of the fascicles.
Female genital pore between the 7th and 8th somites ( 9 , Fig.
119).

Abdominal somites notably longer than the thoracic, beginning at
the 10th, which differs but slightly from the 8th and 9th of the
thorax ; increase caudad in length and number of rings. Uncini-
gerous tori placed near the posterior boundary of each segment ;
the ventral the first to appear, and larger than the dorsal through-
out anterior region of abdomen.

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416 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

Capillary setae (Fig. 120) alone present in first seven segments;
obtusely angled, with striated limb on convex border, arranged in
dorsal and ventral widely separated fascicles ; persist in dorsal fasci-
cles as far as 10th somite ; beyond this point replaced entirely by
nncini.

Uncini (Fig 121 ) begin in ventral fascicles at 8th somite ; in dor-
sal fascicles at the 10th ; hooded, with four minute teeth above ros-
trum; shaft strongly geniculate.

Length of 39 anterior somites, 36 mm. ; greatest transverse
diameter of thorax, 1.5 mm.

This species is represented in the collection by a single incom-
plete specimen, lacking the posterior portion. It is an immature
female, and was collected by Miss Robertson at Port Orchard, July
2, 1898. The dorsal setae of the 8th and 9th somites appear to be
entirely wanting.

CULORAEHIDAE.

32. Trophonia papillata sp. nov. PI. 12, figs. 122, 123.

Form rather long and slender, slightly tapered, thickest anteri-
orly, abruptly diminishing toward mouth ; subcyhndrical ; inters^-
mental constrictions distinct, but not noticeably deepened caudad ;
entire surface papillate, slightly rough to the touch, but without
adherent sand-grains; dorso-ventral differentiation slight, mainly
expressed in differences between dorsal and ventral setae, and
in the closer papillation of dorsum.

Oral tentacles (Fig. 122) eight, of moderate length, pointed at
tip; palpi (/>.) thicker and blunter, grooved on ventral aspect,
distinctly constricted at regular intervals.

iSetae of second, third, and fourth somites, both dorsal and
ventral, elongated, flexible, capillary bristles, forwardly directed
(Fig. 122), exhibiting the usual transversely-striate structure.
Dorsal setae of remaining somites, capillary, three or four to each
fascicle ; ventral setae (uncini), to the same number; stouter than
dorsal setae, blackish, hooked (Fig. 123).

Somites of only complete specimen, 89 in number; those in the
anterior region twice as broad as long; posteriorly, length and
breadth gradually becoming equal.

Lengthy 88 ram. ; greatest transverse diameter, 4 mm.

Two specimens, one incomplete, were collected at Port Orchard,
July 2, 1898, by Miss Robertson.

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JOHNSON; POLYCHAETA OF PUGET SOUND REGION. 417

33. naboUigera infandibnlaris ep. no v. PL 12, figs. 124-
127.

Form (Fig. 124) rather stout, squarish, dorsal aspect more
flattened than the ventral; enclosed in clear mucus; tapered in
posterior third to a minute caudal extremity ; oral region (2d
somite?) flared, with an almost complete circle of setae on the
margin, formed by two broad fan-shaped fascicles ; intersegmental
constrictions deep; parapodia (Figs. 124, 126) well developed,
distinctly biramous, on every somite from the third.

PeriatomiHm within the oral funnel ; bears numerous slender
tentacles and two stout, lobulated palpi (/>.).

Dorsal setae (Figs. 125, 126) longest ^d most numerous on
second somite, where they form the funnel ; on the other somites,
more slender, delicately curved, completely imbedded in the jelly,
transversely stnate. Ventral setae (Fig. 126) begin on third somite,
one or two in each ventral ramus, in form of long hooks, trans-
versely striate, blackish towards tip. Numerous sensory papillae
(Fig. 127) are borne at the tips of long varicose pedicels.

Somites in four specimens are 42, 50, 56, and 71, increasing in
number with size of animal. In contraction, somites are at least
four times as broad as long.

Length of specimen with 56 somites (about average size), 55
mm. ; greatest transverse diameter, 5 mm.

According to the statements of Harrington and Griflin ('97, p.
162), this species is enormously abundant in Scow Bay, where it
covers the muddy bottom over an area about half an acre in extent.
The depth given for one lot of specimens is six fathoms. It does
not appear in any of the littoral collections from the region of
Seattle.

The extraordinary elongation of the dermal sensory papillae in
species of this genus is well exemplified in the present form. The
thick coating of mucus which envelops every part of the animal
except the anterior and posterior extremities (Fig. 124) (the funnel
formed by the broad flabellate oral tentacles makes a passage to the
mouth) apparently renders necessary this method of putting the
animal in communication with the outer world.

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418 PROCEEDINGS: BOSTON SOCIETY NATURAL mSTORT.

Stebnaspidae.

34. Stamaspis fossor (?) Stimpson.

Three specimeDs, apparently of • this species, were collected at
Victoria, Vancouver Island, by Prof. W. A. Herdman. The speci-
mens are not sufficiently well preserved to admit of thorough and
critical study ; but comparison of the ventral shields with those of
a specimen of /S. /ossor from the Atlantic coast, and also with
Marenzeller's ('90) figures, makes it reasonably certain that the above
identification is correct. Specimens from Puget Sound collected by
the naturalists of the Northwest Boundary Commission were doubt-
fully described by Stimpson (*64) as a new species, which he named
Sternaspia nffinis.

Maldanidab.

85. Clymenella ^ rnbrocincta sp. nov. PI. 13, figs. 128-133.

Form considerably elongated, cylindrical, narrowed in region of
third and fourth somites, gradually enlarging to maximum diam-
eter in region of somites 10-12, thence narrowing to somites 15-17,
which are the longest and slenderest of the body ; the remaining
somites (18-22) slightly thicker and progressively shorter; the
2l8t the shortest of the body.

Cephalic plate (Figs. 128, 129) oval, concave on dorsal side,
nearly bisected longitudinally by a median ridge continued back from
the ovate palpode ; a distinct raised margin, and well-developed
longitudinal nuchal organs (Fig. 129, n, o.). Mouth with thick-
ened corrugated lower lip and crescentic outline.

Peristomiwa achaetous ; capillary setae and uncini begin on sec-
ond somite (Fig. 128), the latter with scarcely perceptible tori at
first ; tori become distinct in fourth somite ; dorsal setae from wart-
like papillae.

tSohiites gradually increasing in length from fifth onward ; fifth
to eighth marked with a broad indian-red band back of the setae
and uncini, accentuated by a narrower whitish band in front.

» Axiothea and Clymeyiella cover species too nearly alike to require generic distinc-
tion. As recently pointed out by Verrill ( :00, p. 657). the name Axiothea being preoc-
cupied, Clymenella {semu ext.) should cover the species formerly included under
Axiothea.

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JOHNSON: POLYCHAETA OF PUGET SOUND REGION. * 419

Tori much enlarged in somites 12-21. Last 8 somites (including
pjgidium) achaetous; pygidium funnel-shaped with eirrose border.
Perianal cirri (Fig. 130) 18-30 in number, alternately long and
short ; the mid-ventral one much elongated, and containing prolonga-
tion of the ventral nerve-cord ; tips of all the longer cirri recurved.
Anal rosette with alternate sectors raised, corresponding in position
with the longer cirri.

CapiUary setae of two forms, bordered and serrated, the latter
much the more slender (Fig. 133). Both kinds in same fascicle;
begin at second somite. Uncini (Figs. 131, 132) with five or six teeth,
including rostrum, graduated in size ; bristles at base of rostrum lat-
eral i^ position.

Length of large specimen, 162 mm.; greatest transverse diam-
eter, 3.5 mm.

This fine species comes near Axiothea ccUenata Malmgren, a form
of wide distribution in the North Atlantic and Arctic, and reported
from Bering Sea by Marenzeller ('90). The present species, how-
ever, differs from it in the form of the serrated setae and of the
uncini. In A, catenata there are four preanal achaetous somites,
in the present species only two ; C, ruhrocincta has 22 segments ;
A, catenata, 24.

The prfesent species was collected both by the Columbia Univer-
sity Expedition and by Miss Robertson. I have found it in
abundance at the entrance of Tomales Bay, and at San Pedro,
California. It forms a tube of coarse sand, which is lined by a
peculiarly tough, opaque, whitish membrane.

36. Nicomache personata sp. nov. PI. 13, figs. 134-139.

Size small ; 25 somites, of which all except the peristoraium and
the last two are setigerous; no cephalic plate; prostomium and
peristoraium united to form a hood-shaped head (Fig. 134);
mouth large, overhung by projecting upper lip ; a distinct crease in
peristoraium back of mouth.

Somites increasing in length back of second ; longest from the 8th
to the 17th; those at the end very short; pygidium (Fig. 135)
funnel-shaped, with a zone of 16-18 cirri on margin, quite uniform
in length. Dorsal and lateral surfaces of head and first four or five
somites beautifully mottled with chocolate-brown; ground-color,
white ; somites 2-5 with contrasting white and brown bands ; on the
head these bands have the aspect of a grotesque face.

Setae of five kinds: (1) capillary double- bordered setae in all

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420 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

somites from second to twenty-third; (2) smaller serrated setae
together with (1) in dorsal rami; (8) ventral setae in first three
setigerous somites in form of a stont, acate spine (Figs. 134, 136) ;
(4 ) very long, filamentous, spiral setae in dorsal rami beginning at
7th to 11th somites and continuing to 23d somite (Fig. 138); (5)
uncini {Fig. 137) of the usual form in this family, in all setigerous
somites back of the 4th.

Length of much-contracted specimen (impossible to measure
accurately on account of twists and flexures), 50 mm.; diameter
through thickest portion, largest specimen, 1.75 mm.

The only specimens of this odd little species were collected by
Hiss Robertson at Alki Point, Aug. 3, 1898. The specie^ bears
a close superficial resemblance to Xicmnache lumbricalis (Sars)
Malmgren, but has one achaetous somite in front of the pygidium
instead of two.

The long, silky, spiral filaments shown in Fig. 138 have not, I
believe, been, hitherto described in any Maldanid. Whether they
are permanent structures peculiar to this species, or " nuptial setae '*
(Pubitfttsborsten) which develop only at sexual maturity, is an
interesting question. Filamentous nuptial setae of a similar form
have been described by Michaelsen ('92, p. 6) in a Polynoid
(Drieschia pelagica),

Ammocharidae.

37. Ammochares occidentalis sp. nov. PI. 14, figs. 140-142.

Form cylindrical, tapering towards posterior end ; 23 (?) somites,
of which 20 are setigerous.

Peristoinium produced into ten thick, bluntly-ramose '* tentacles"
(Fig. 140) which rise to a level, giving anterior end of body a trun-
cated appearance. Capillary setae in fascicles placed high on dor-
sal surface (Fig. 142), very slender, acutely pointed, silvery by re-
flected light, serrated. Uncini very minute, two-hooked (Figs. 141
a, ^), lG-18 horizontal rows in each band (6th somite), extend-
ing three quarters of the way around the body ; begin at 4th somite.

Second and third so)nites (Fig. 140) very short, third twice the
length of the second, both with rudimentary parapodia (?) . Somites
4, 5, 6, 7 the longest of the body ; from the 7th diminishing gradu-
ally to end of series.

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JOHNSON: POLTCHAETA OF PUGET SOUND REGION. 421

Length, 22.5 mm. ; greatest transverse diameter, 1 mm.

No species of Ammochar€8 has hitherto been reported as having
chelate or bifid uncini. This character in fact has been considered
so diagnostic of the allied genus Myriochele that M'Intosh did not
hesitate to describe a species in the " Challenger '' collection as My-
riochele pacifica from a fragmentary specimen lacking both anterior
and posterior extremities — basing his diagnosis entirely on the
structure of the uncini.

The two specimens upon which the foregoing description is based
were collected by Miss Robertson at Port Orchard, July 2, 1898.
£ach was enclosed in a tube composed of sand-grains and minute
particles of shell. The color of the formalin preserved specimens is
nearly black.

Arenicolidae.

38. Arenicola claparedei Levinsen. PI. 14, figs. 143, 144.

In the excellent memoir of Gamble and Ashworth (:00) upon
the Arenicolidae, the Mediterranean species, originally described
by Clapar^de ('70, p. 300) as Arenicola marina, but afterwards
erected as a separate species by Levinsen ('83, p. 137, footnote)
under the name of A. claparedei, is attributed to the Pacific coast.
After a careful examination of the Puget Sound Arenicolae at my
disposal, and comparison with specimens of A, claparedei from
Naples, I am convinced that Messrs. Gamble and Ashworth are cor-
rect in their determination. The only notable points of difference
between the Puget Sound specimens and those from Naples are the
vastly greater size — at least eight times as great — of the former,
and the smaller number of oesophageal coeca or "pouches" in the
latter. Of the four specimens of A, clffjyaredei from Naples which
I have examined, three have four pairs of pouches, and one only
three pairs ; whereas, out of eight specimens from Puget Sound in
four there are six pairs, in two fifteen pairs, in one sixteen pairs ; and in
one there are sixteen coeca on the right and eighteen on the left !

In the Arenicolidae there are as a rule only two oesophageal
coeca or " pouches," but in A. cla2xtredei they are not only numer-
ous (as many as 32 in one instance, Fig. 144) but highly variable as
to number and arrangement, and probably even differ as to function,
if it is permissible to draw such an inference from the great size and
thin-walled character of the most anterior pair (Figs. 143, 144).

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422 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

Ab, howerer, the whole qaesdon of the stmcture and functioii of
the oesophageal pouches of Arenicola is still unsolved, this problem
must be deferred for the present.

The discovery of this Mediterranean form upon the Padfic coast
of North America, and at no intermediate point, is certainly one of
the most remarkable cases of discontinuous geographical distributioa
ever recorded.

With exception of a single specimen collected by Miss Robertson,
I am indebted to Dr. C. M. Child of the University of Chicago for
all the Puget Sound Arenicolcte I have. The specimens were col-
lected by the Columbia University Expedition — exact locality not
stated.

While, as von Marenzeller has suggested, it is probable that
Murdoch's ('84) ArenicoUi glacialis from arctic Alaska is none
other than the circumboreal and widely-ranging A. marina, it is not
at all certain that the latter species occurs so far south as Puget
Sound, although it has been reported from Vancouver Island by
von Marenzeller ('87).

CiRBATULlDAE.

39. Cirratalns cingolatus sp. nov. PI. 14, figs. 145-148.

Form stout, size moderate, tapered at both extremities, decidedly
flattened on ventral aspect; dorsum rounded; seven anterior
somites without setae or cirri ; two clusters of 17-18 tentacular fila-
ments (Fig. 145, t.f.) each, on dorsal aspect of 8th (the first setig-
erous) somite; when removed, an oval transverse scar is left; a
series of similar cirri along each side, in the anterior and middle
portions of body inserted low (Fig. 146), gradually rising to a higher
level in the posterior region.

SoNuteH very short, three-ringed above the dorsal setae, the
middle ring raised welt-like above the level (Fig. 146).

Prostortiimn (Fig. 145) acute, concave on ventral side towards
mouth, which usually exhibits a partially everted pharjTix; eye-
spots five or six, either in a group or transverse row.

Pumpodhi slightly developed (Figs. 145, 146), both dorsal and
ventral rami with slender, serrated setae (Fig. 147) ; these alone
are present for 30 setigerous somites back of the head ; the uncini
appear in the ventral rami (Fig. 148) at this point, and in the
dorsal rami a few somites caudad ; they continue to end of series.

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JOHNSON : POLYCHAETA OF PUGET SOUND REGION. 423

Cirri, especially those of the first cirriferous somite ("tentacular
filaments^') of great length, frequently forming a tangled mass
nearly as long as the entire body. Anus in pygidium, minute.

Length of large specimen, 100 mm. ; greatest transverse diam-
eter, 5 mm.

There are several specimens in the Columbia University collec-
tion, obtained " between tides." I have not found this species on
the California coast.

40. Cirratulns robnstiui sp. nov. PI. 14, figs. 149, 150.
Form short and thick ; ventral aspect broadly flattened, concave

anteriorly and posteriorly ; dorsum rounded, tapered about equally
towards both ends; 112 somites in one specimen, not dimded into
rings (Fig. 150).

Prostomitim (Fig. 149) shorter and thicker than in preceding
species ; eye-spots in two oblique rows. First three somites achae-
tous, fully twice the length of the rest ; fourth with two clusters of
tentacular fllaments and with dorsal and ventral setae (Fig. 149, t. /.).

Parapodia (Figs. 149, 150) slightly developed; anteriorly with
capillary setae alone; at the 19th or 20th somite the ventral uncini
begin, and two or three somites further back, the dorsal; cirri
inserted low in anterior somites (Fig. 149), gradually rising to mid-
lateral line (Fig. 150) towards middle of length, then in posterior
region gradually approaching the dorsal setae. Anus terminal.

Lengthy 59 mm. ; greatest transverse diameter, 5 mm.

Only two mature and one yoiung specimen of this species are
available for description. One adult specimen was obtained at
Neah Bay by the Columbia University Expedition ; the other at
Port Orchard by Miss Robertson. Both lack the cirri. The setae
and uncini of this species are practically identical with those of
Cirratidus clngulatus,

Amphictenidae.

41. Fectinaria brevicoma sp. nov. PI. 15, figs. 151-156.
Fortn conical, gradually widening towards anterior end ; cephalic

disc nearly flat, plane, its edge entire ; scapha broadly ovate, not
wider transversely than posterior end of thorax. Total number of
somites, about 27 ; 21 in thorax (of which 17 are setigerous, and 13
are uncinigerous), and 6 (?) in scapha.

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424 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

Pro8tom\um expanded into a large velnm (Fig. 151) with 33 fim-
briae on edge.

Periatornium with a pair of subulate, mpniliform cirri and numer-
ous club-shaped tentacles (much contracted in preserved specimens).
Paleolae short, stout, arranged in two distinct groups, 10-12 in each ;
highly iridescent, either blunt or acutely pointed (lateral ones),
strongly sloped towards ventral aspect. Second somite with a pair
of cirri in all respects like the peristomial.

Branchiae pectinate, borne on third and fourth somites.

Setae broadly limbed and twisted near tip (Fig. 152) with serra-
tions beyond the twist, or straight and without serrations (Fig. 153) .
The latter are not so numerous. They diminish in size towards
anterior and posterior extremities of thorax. Uncini with 4 teeth
(Fig. 154) or occasionally five (Fig. 155) ; the latter begin about
the 11th somite; none were found in front of this. Spines on
scapha (Fig. 156) with a stout, laterally-bent hook at tip.

Tube composed of coarse sand-grains, curved.

Length of largest specimen, 28 mm. ; diameter of disc, 5 mm.

Several specimens were dredged by the Columbia University
Expedition at a depth of 10 fathoms.

This species comes nearest to P, {Cistenidea) granulata
(Malmgren), which has been found in Bering Sea (Marenzeller,
*90), and was collected at Kadiak by Mr. Cloudsley Rutter, differ-
ing from it only in the shortness of the paleolae and in the form of
the setae and uncini. Upon examination of more abundant ma-
terial, this form may prove to be identical with grannlata^ which is a
wide-ranging, circumboreal 8i)ecies, and may therefore prove to be
variable. While the uncini afford, on the whole, the surest diagnos-
tic characters, they should be used with caution, as their variability
in the present species clearlv indicates.

Ampharetidae.

42. Sabellides anops sp. nov. PI. 15, figs. 157-161. PI.
16, figs. 162-163.

Form stout, curv ed, thickest about midway of thorax, abdomen
rather rapidly tapered, convex on dorsal, flat on ventral aspect
(Fig. 157). Thirty to thirty-one somites, sixteen in thorax, four-
teen to fifteen in abdomen ; fourteen setigerous somites in thorax ;

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JOHNSON: POLYCHAETA OF PUGET SOUND REGION. 425

setae begin at third somite and cease at sixteenth. Uncini begin
at sixth somite (the third setigerous somite), and extend to the
pjgidium. Dorsal rami of all abdominal parapodia destitute of
setae and uncini.

Prostoinium (Figs. 158, 162, 163) broadly truncated with two
anterolateral lobes ; no eyes ; a pair of transverse slits containing
the nuchal organs at its base.

Peristamuim (Figs. 158, 162, 1) forming, with second somite, a
region destitute of parapodia and branchiae. The club-shaped
tentacles, about fifteen in number, arise from the inner border of the
peristomium ; they are destitute of papillae, but in contracted state
are wrinkled (Fig. 163).

Branchiae^ four on each side, smooth, terete, subulate, arise from
dorsal aspect of the third, fourth, and fifth somites ; in length about
twice the diameter of the body (Figs. 158, 162).

Ventral rami or tori (Fig. 157) spatulate, increasing from the
fom'th somite to the sixteenth, thence diminishing to end of body.
Each bears a single row of pectinate, six-toothed uncini (Fig. 161).
Dorsal thoracic rami contain fascicles of single and double-
bordered, straight and slightly-curved setae (Figs. 159, 160);
dorsal rami achaetous throughout the abdomen.

Length of largest specimen, 27 mm. ; greatest transverse diameter
of same, 5 mm.

Several specimens are in the Columbia University collection,
without data as to depth and locality. No tubes were preserved.

This species comes close to SabeUidea {Amage) auricula (Malm-
gren) but differs from it in having longer branchiae, attached to
three somites, shorter tentacles, and differently shaped uncini.
SdbeUides auricida^ however, has been reported from Japan by
Marenzeller ('85), and its occurrence in any part of» the North
Pacific would therefore not be surprising.

I follow Theel ('79) in discarding Malmgren's genus Amage, as
not being sufliciently distinct from Sabellides,

Terebellidae.

43. Amphitrite robusta sp. nov. PI. 16, figs. 164-168.
Form short, robust, thickest anteriorly in region between 5th and
12th somites, gradually and almost uniformly tapering thence to

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426 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

posterior end. Total number of somites, 63-90 ; in thorax, 20, of
which 17 (4th to the 20th inclusive) are setigerous. Uncini begin
at 5th somite and extend to the penultimate. Uncini nniseriai
anterior to the 11th somite, and posterior to the 20th; biserial on
thoracic somites 11-20 (Fig. 167).

Prostomiiim with dorsal ridge and small lateral lobes formed by
its continuation to the sides; no eyes. Peristomium bearing a
semicircle of tentacles, rather thick in formalin specimens, decidedly
grooved, spirally coiled in contracted state, and about one half the
length of the thorax. Peristomium forms a thick prominent ventral
lip, opposible to thin ventral edge of prostomium.

Bratichiae (Fig. 164), three pairs, on somites 2-4, densely
ramose, di- and tri-chotomously branched ; the main stems short and
thick, ultimate branches subulate, slightly moniliform. Branchiae
all nearly alike in form and size ; anterior pair slightly the largest ;
all variable as to size and amount of branching.

S€ti(feron8 lobes moderate, increasing in size from the first to the
seventeenth (on 20th somite). Uncinigerous tori of nearly uniform
length from 5th to 18th somites; thence gradually diminishing to
end of body.

Setae with striated limb on each side and curved, serrated tip
(Fig. 165). Uncini avicular, with from 5 to 7 rows of teeth above
the rostrum (Figs. 166-168).

Xejt/iridi'al 2)apiUae twelve psirs^ on somites 4 to 15; first pair
considerably the largest ; the rest of nearly uniform size ; placed
ventrad, and a little posterior to setigerous lobes.

Lenffth of larger specimen, 75 mm. ; greatest transverse diameter
of thorax, 11 mm. ; average transverse diameter of abdomen, 4 mm.

Several specimeim, collected both by the Columbia University
Expedition and by MiHs Robertson. The species is doubtless com-
mon. No tubes were preserved. The absence of eyes in this
species and in the preceding is remarkable. They are evidently
wholly lacking, as I was unable to find them even in serial sections.

44. Amphitrite spiralis sp. nov. PI. 16, figs. 169-171c.

Form greatly elongated, abdomen slender, terete, and spirally
coiled when out of the tube ; dorsal aspect of thorax high-arched ;
ventral slightly convex. This condition is enhanced in anterior
portion of abdomen,, where the somites are decidedly thicker on
the dorsal than on the ventral aspect, producing thereby the spiral

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JOHNSON : POLYCHAETA OF PUGET SOUND BEGION. 427

coiling. Number of somites approximately 170 ; those of posterior
portion of abdomen two-ringed. Thoracic somites, 43, of which 41
are setigerous and 39 uncinigerous.

Prostomimn with dorsal crescentic groove separating a dorsal
ridge from the ventral flap ; no eyes visible on surface.

Peristomiiim with deep ventral groove, and dorsal semicircle of
cirri.

Branchiae two pairs, on third and fourth somites ; moderate ;
anterior pair considerably the larger and m6re richly branched;
main branches arising near the base ; beyond the first ramification,
the branching is dichotomous (Fig. 169).

Setir/erous lobes begin on fourth somite, increase slightly in size
towards middle of thorax, then diminish ; the last few pairs very
small. Setae of form usual in this genus (Fig. 170).

Uncinigerous tori begin at fifth somite; first six pairs shorter
than the rest ; gradually increasing in length to the seventh where
they attain the maximum length, and retain it to the 16th or 17th, ^t
which point they gradually diminish, becoming flattened and almost
indistinguishable on the abdomen. Uncini small, avicular, with five
rows of teeth above the rostrum (Fig. 171 c). They are uniserial
on somites 5 to 10, and on 35 to end of series ; biserial, on somites
11 to 34.

Lengthy 160 mm. ; greatest transverse diameter of thorax, 5 mm. ;
average of abdomen, 2 mm.; dorso-ventral diameter of thorax, 4.5 mm.

A single individual of this species was collected by Miss Robert-
son near Seattle.

45. Lanice heterobranchia sp. nov. PI. 17, figs. 172-174.

Thorax of 20 somites, 17 (?) setigerous.

Prostonnuni of usual form ; no eyes.

Branchiae three pairs, borne on somites 2-4; first pair (Fig. 172)
much the largest and longest, with elongated main stem dendrit-
ically branched ; ramifying branches very compact. Gills of second
and third pairs short, without main stem.

Setae (Fig. 173) with striated limb on each side ; tip entire, from
fourth (?) somite onward. Uncini from fifth somite, uniserial, alter-
nating ("rangee alterne," Clapardde), avicular, with three teeth in
front of beak (Fig. 174).

A single specimen in the Columbia University collection, too
imperfect for complete description. The worm was enclosed in a
mud tube. The difference in the size of the branchiae of different
pairs is the most striking character.

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428 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

46. Tholopns crispns sp. nov. PL 17, figs. 175-178^.

Form rather stout, not greatly tapered posteriorly ; thorax pass-
ing gradually into abdomen ; tapered considerably towarcis prosto-
mium from tenth somite. Number of somites 88-147.

Prostomium with ample dorsal flap, transversely corrugated on
dorsal surface ; no eyes.

Peristotnium with circlet of strongly grooved tentacles.

Branchiae (Fig. 175) three pairs, branching from the base in
numerous, slender, spirally curled filaments ; borne on somites 2-4.

Sttae, begin at third somite and extend to penultimate in young
specimens ; to fourteenth from pygidium in older ones ; with stri-
ated limb on each side; sometimes slightly bent (Figs. 176, 177).

Uncinigerous tori begin at fourth somite; nncini absent from
extreme end of body. Uncini (Figs. 178 a, ^) single-ranked from
fourth to seventh somites, inclusive, and gradually returning to this
condition towards end of series; in flattened rings ^m eighth
somite (^^rang6e parabolique," Clapar^e) onward. Tori attain
their greatest length between the twelfth and twenty-fourth
somites ; thence diminish very gradually to end of body ; those of
the abdomen rounded and wart-like.

Length of large female specimen, 270 mm. ; greatest transverse
diameter (at sixteenth somite), 13 mm.; dorso-ventral diameter,
12 mm.

This flne Terebellid is represented in the Columbia University
collection by a single large female turgid with eggs. It occurs on
the California coast as far south at least as San Francisco, and is
abundant at Bolinas, Marin County. Its tube is formed of coarse
sand or gravel. It frequently harbors commensal individuals of
Pohjnoe insignis^ and northward, probably also Hamiothoe tuta (see
p. 394).

Sabellidae.

47. Bispira polymorpha sp. nov. PI. 17, figs. 179-183. Fl.

18, ?i^i^, 184, 185.

Fonn nearly terete, dorsiira flattened in adult specimens; in
young specimens, form slender; tapered gradually at posterior end
to a minute pygidium; anus terminal. Somites, 170 or more.

Thorax (Fig. 179) of nine somites, one sixteenth to one ninth of
entire length (exclusive of branchiae) according to degree of con-
traction and probably also the age of the specimen.

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JOHNSON : POLYCHAETA OF PUGET SOUND REGION. 429

Branchiae (Fig. 179) about as long as thorax, dichotomously
branched twice or thrice in some specimens, in others unbranched ;
pinnate, radioles slender; 16-30 branchiae on each side, forming a
spiral of 2-3 turns ; each rachis with 2-10 black eye-spots (Fig. 184).

Tentacles flattened, lanceolate, about one fifth the length of
branchiae.

Fecal groove extending forward along mid-dorsal line of abdomen
to thorax ; at posterior boundary of thorax passing on the left side to
mid-ventral line of same, and thence to oral region.

Peristoniium with raised anterior border or " collarette," deeply
notched in mid-dorsal line, and produced into two pointed processes
adjoining the ventral sulcus.

Thoracic setae (Figs. 180, 181) begin on second somite; of two
forms, winged-capillary and mucronate-spatulate ; the latter more
numerous, forming a close series towards the torus.

Uncinigerous tori of thorax dorsal to setigerous tubercles ; begin
on third somite ; separated by full width of dorsum ; uncini biserial,
of two sorts (Fig. 182), avicular and dilated-cuspidate ; both kinds
with long manubria ; the points directed cephalad. Abdominal setae
all of one kind (Fig. 185) arising from smaller tubercles, which are
placed dorsad to the uncinigerous tori. Uncini (Fig. 183) uniserial,
all avicular, with a shorter manubrium than the thoracic uncini ;
rostra directed anteriorly.

Tube cartilaginous, translucent, adherent to rocks, piles, etc.

Length of average specimen (exclusive of branchiae), 96 mm.;
greatest transverse diameter, 6 mm. Greatest transverse diameter
of largest specimen at hand, 12 mm.

Numerous specimens from the Puget Sound region, collected at
Neah Bay and in the Port Townsend district by the Columbia Uni-
versity Expedition, and at Alki Point and Port Orchard by Miss
Robertson. It occurs also on the California coast as far south as
Pacific Grove.

This species is remarkable for the highly variable aspect which it
presents, owing to the diverse coloration of the branchiae and the
differences of shape caused by different states of contraction in
which it has been killed. If killed within the tube, it is almost per-
fectly cylindrical and often of great length, owing to the impossibil-
ity of expansion within the rigid and tightly fitting tube. The
longest specimen thus killed (posterior somites lacking) measures
not less than 150 mm. in length and only 5 mm. in greatest trans-

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430 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

verse diameter. Even more striking, although not always percep-
tible to the naked eye, is the variable character of the branchiae,
which are in some specimens dichotoraously branched, and in others
entirely unbranched.

The coloration of the branchiae shows two distinct phases — pur-
ple or wine-color and whitish or tawny. Either color may be present
to the exclusion of the other, or the two may be in alternate, trans-
verse bands. The eye-8[)ots may be few or many, but I have found
no specimen without them. Although a lens is absent, the eye-spot
j)roduces a wart-like elevation of the cuticula which covers it (Fig.
184). The eye-spots are of various sizes, the largest being over
twice the diameter of the smallest. They are scattered at irr^ular
intervals along the rachis, but are more numerous towards the base
than towards the tip. None are found near the tip and very rarely
any on the branches.

The tube is adhesive, and is usually aUbced by its lower extremity
or by its side to a stone or pile. At Pacific Grove, the species attains
a large size and is abundant on rocky bottoms from low-water mark
to a de})th of several fathoms.

Megachone gen. nov.

Form terete; no ventral shields; collarette liaring, interrupted
only at ventral notch ; branchiae connected by a web ; no spatulate
thoracic setae ; thoracic uncini with long manubrium, of one kind
only ; abdominal uncini short avicular plates ; no ventral fissure at
posterior end.

48. Megachone aurantiaca sp. nov. PI. 18, figs. 18G-192.

Form subcylindrical, spiral in contraction, thickest in posterior
jwrtion ; anterior end truncated; collarette broad (Figs. 186, 187) ;
body narrowed in region of thorax and enlarged in abdomen ; pos-
terior extremity abruptly tapered to a minute tip, which is curved
dorsad ; anus terminal. Somites 75 in number.

Branchine 20 on each side, unbranched (Figs. 187, 188), closely
appressed, tips pointing ventrad ; radioles invisible until branchiae
are raised.

Th(jr<ix (Fig. 186) of eight somites ; peristomium, with collarette,
one and one half times the length of the other thoracic somites;
bears a small tuft of setae, but no uncini ; a slight indication of two
rings ; the biannulate condition more strongly marked in the other
thoracic and abdominal somites.

Fecal groom extending forward along mid-dorsal line to ninth

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JOHNSON: POLYCHAETA OF PUGKT SOUND REGION. 4^31

somite, there passing on the left side to mid-ventral line, and thence
to oral notch. Setigerous tubercles small.

Capillary seUie (Fig. 189) of thorax single or double-bordered ;
uncinigerous tori (Fig. 186) short, small ; uncini avicular, with very
long manubria, uniserial, or incompletely biserial (Fig. 190); setae
of the abdomen very similar to those of thorax ; uncini of abdomen
(Figs. 191, 192) with much elongated rostra.

Lenffth^ 87 mm. ; transverse diameter of collar, 5 mm. ; greatest
transverse diameter of abdomen, 4.5 mm.

This species is represented by a single specimen, collected by Miss
Hobeitson at Port Orchard, on July 4, 1898. The specimen was
preserved in formalin and the color when first received (no doubt
nearly that of the living worm) was a bright orange. The worm is
undoubtedly a tube-dweller, but there are no data concerning a tube
or the precise habitat.

As suggested to me by Professor Verrill, the branchiae of this
specimen are in all probability in process of regeneration.

EUIOGRAPIIIDA^.

49. M3rxicola pacifica sp. nov. PI. 19, figs. 193-198.

Body fusiform, terete ; tapered slightly towards anterior, much
more, and gradually, towards posterior extremity ; somites 67-97 in
numl>er, short, biannulate; no collarette; two fleshy processes
(tentacles?) on first somite, adjacent to mouth; thorax hardly dis-
tinguishable from abdomen and composed of nine somites.

Branclnae (Fig. 193) 14 on each side, connected by a web (w,)
as far as radioles extend, /*. e,^ within 3 mm. of tip ; radioles slender,
biserial; ti{)s of branchiae spirally coiled in retraction.

Thoracic setae of two forms: (1) double-bordered capillary
(Fig. 194), very slender, often twisted; and (2) blunt, spinous
setae (Fig. 195) with conical tips, much fewer in number; both
kinds occur together in tufted fascicles. Uncini of thorax with
long manubria (Fig. 197), strong rostra, and minute teeth ; uniserial.
Abdominal setae (Fig. 195) slender, minute, very broadly double-
bordered ; alxlominal uncini (Fig. 198) very minute, avicular, 2-3
teeth above rostrum; arranged in tori which nearly encircle the
body, being interrupted only by the setigerous papillae and a
median stripe on doi*sum ; uniserial.

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432 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

Length of large specimen (not including branchiae), 60 mm.;
greatest transverse diameter of same, 9 mm. ; length of retracted
branchiae, 21 mm.

This species is represented by two female specimens, one col-
lected by Miss Robertson at Port Orchard in June, 1899, the other
by Professor Ritter at Pleasant Beach in May of the same year.
Although exhibiting great difference as to size — one being more
than twice the length of the other — they both contain nearly ripe
ova, indicating sexual maturity.

The transparent mucous envelope so characteristic of this genus
was preserved with both specimens.

Sebpulidae.

oO. Serpula columbiana sp. nov. PI. 19, figs. 199-204.

Form subterete; somewhat flattened dorso-ventrally ; gradually
tapered towanis posterior extremity; fecal groove distinct, dorsal
as far as thorax, there branching and passing to the ventral side
under the thoracic membrane.

Thorax with seven setigerous somites, well-developed collarette,
and thoracijB membrane, the latter reaching to the tips of the setae.
Abdominal somites, 250 or more.

Branchial filaments 54 on each side, arranged in two spirals
ascending from the ventral edges, each making a complete turn,
then extending mesad and meeting each other just over the ibonth.
Branchiae whitish, beautifully banded with scarlet or crimson ; distri-
bution of color variable but usually a broad red band at or near
base, followed by two narrow bands, and branchiae broadly tipped
with the same. Operculum (Fig. 199) on right side, its mate on
the left very short and rudimentary ; funnel-shaped, with about 100
ribs which form a notched border; deep, funnel-shaped cavity;
pedicle geniculate just below the operculum.

Bayonet-setue of first setigerous somite as in Fig. 200 ; the other
thoracic setae broadly striate-bordered (Fig. 201). Uncini of
thorax and abdomen similar in shape, 6-S toothed ; the tip of
largest tooth often turned outward (Figs. 202, 203). Chisel-shaped
setae of abdomen as shown in Fig. 204.

Tube white, calcareous, more or less coiled ; anterior portion of
old tubes often free from the substratum to which tube is attached.

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JOHNSON : POLYCHAETA OF PUGET SOUND REGION. 433

Length of large specimen, 55 mm. ; greatest transverse diameter
of thorax, 7 mm. ; of abdomen, 6 mm.

This beautiful Serpnlid is abundant in Puget Sound. Harrington
and Griffin ('97, p. 103) mention a handsome Serpula^ probably
this species, which forms " hard white tubes " on the rocky bottom
of Hood's Canal ; and also as " whitening the rocks with its calca-
reous tubes," along the beaches near Port Townsend. It was col-
lected at Alki Point by Miss Robertson. It occurs also on the Cal-
ifornia coast at Bolinas (Duxbury Reef) , and at Lime Point and
Point Cavallo, on the northern shore of the Golden Gate. Its
favorite habitat is the under side of a stone where the water flows
freely. If not in a tide-pool, it is near extreme low-water mark.
The uncini show considerable variation on the same individual,
and even on the same torus. The number of teeth ranges from six
to eight ; the upper border is high-arched or nearly straight. The
tip of the large tooth may or may not be recurved.

The coloration of the branchiae is also variable, both as to tint
and distribution. It is either scarlet or damask-red, *and it may in-
volve nearly the whole of the branchiae and operculum, or may be
more restricted, so that the white predominates. If formalin speci-
mens are not exposed to direct sunlight the color is retained for
years in almost its original brightness.

This may possibly be identical with Sevpvla jttkesii Baird, de-
scribed by Grube ('77) from North Japan. His description is too
brief, however, to put the question of identity beyond doubt.
51. Serpula zygophora sp. nov. PI. 19, figs. 205-208.
Form nearly cylindrical ; abdomen strongly grooved on dorsal
aspect, the somites marked on each side by transverse ridges ; seven
thoracic setigerous somites.

Branchiae spiral, thirty filaments on each side; carmine-red at
base, and broadly barred with the same ; operculum (Fig. 205) on
right side, funnel-shaped, moderately cupped, 26-ribbed, the ribs
extending to the center of the concavity ; base yoke-shaped ; pedi-
cle long and curved, geniculate just below the operculum; corre-
sponding filament of left side club-shaped. Operculum and pedicle
variegated with red.

First setigerotis somite with bayonet-setae (Fig. 206). Thoracic
uncini six-toothed (Fig. 207) ; abdominal uncini (Fig. 208) five- or
six-toothed. Chisel-shaped abdominal setae almost identical with
those of Serpula cohunhiana. Transverse diameter of thorax, 3.5
mm. ; of ablomen, 3 mm. Tube lacking.

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434 PROCEEDINGS : BOSTON SOCIETY NATURAL HISTORY.

This species is represented by a single imperfect specimen (lack-
ing posterior portions), collected by Miss Robertson at Alki Point.
As it has never been collected, so far as known, on the California
coast, it may be inferred that its distribution is northward.

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JOHNSON: POLYCHAETA OF PUGET SOUND REGION. 435

LITERATURE.

Andrews, E. A.

*91. The distribution of Magelona. Johns Hopkins univ. circulars, vol.
10, no. 88, p. 96.
Arwidsson, Ivar.

*98. Studien tiberdie familien Glyceridae und Goniadidae. Bergens mu-
seums aarbog, no. 11, pp. 1-69, 4 taf.
Beiiham, W. B. ^ .

*96. The blood of Magelona. Quart, joum. micros. 8ci.,vol. 89, pp. 1-17,
Ipl.
Clapar&de, Ed.

•68-*70. Les Ann61ides Ch^topodes du (Jolfe de Naples. M6m. de la soc.
de phys. et d* hist. nat. de Genfeve, t. 19, pp. 313-584, 16 pis.; lere part.,
pp. 1-225, 15 pis.; 2e part., pp. 365-542.
Darboux, J. G.

'99. Recherches sur les Aphroditiens. Travaux de Tinstitut do zoologie
de Tuniversitfi de Montpellier, etc., nouv. sCr., m6m. no. 6, 276 pp., 83
figs.
Dean, B., N. R. Harrington. G. N. Calkins, & B. B. Griffin.

'97. The Columbia University zoological expedition of 1896, etc. Trans.
N. Y. acad. sciences, vol. 16, pp. 33-42 (with map of Puget Sound).
Ehlers, Ernst.

*64-'68. Die borstenwUrmer (Annelida Chaetopoda), bd. 1, Nereidea, 4to,
748 pp., 24 taf.
Gaml)le, F. W., & J. H. ^Vshworth.

: 00. The anatomy and classification of the Arenicolidae, with some ob-
servations on their post-larval stages. Quart, journ. micros, sci., vol.
43, pp. 419-569, 8 pis.
Grube, Ed.

'51. Middendorffs Reise in den ftussei-sten norden und osten Siberiens,

bd. 2 (Zool.), th. 1. (Annelida.)
'55. Beschreibung neuer oder wenig bekannter Anneliden. Arch. fUr

naturgesch., jahrg. 21 (1855), bd. 1, pp. 81-136, 3 taf.
'1?. Neue Anneliden aus Japan. 55 Jahresber. der Schles. gesellsch. ftir
vaterl. cultur, pp. 104-106.
Harrington, N. R, ' •

'97. On Nereids commensal with Hermit Crabs. Trans. N. Y. acad. sci-
ences, vol. 16, pp. 214-221, 3 pis.
Harrington, N. R., & B. B. Griffin.

'97. Notes upon the distribution and habits of some Puget Sound inver-
tebrates. Trans. N. Y. acad. sciences, vol. 16, pp. 152-165.
Haswell, W. A.

'86. On the structure of the so-called glandular ventricle (drUsenmagen)
of Syllis. Quart, joum. micros, sci., vol. 26, pp. 471-480, 1 pi.

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436 PROCEEDINGS: BOSTON SOCIETY NATURAL HISTORY.

Johnson, H. P.

"97. A preliminary account of the iflarine Annelids of the Pacific coast,
with descriptions of new species. Proc. Cal. acad. sciences, 3d ser.,
(Zoology). voL 1, pp. 153-198, 6 pis.
Kinberg, J. G. H.

'55. Nya slftgten och arter af Annelider. 1. Aphroditea. Ofrereigt

kongl. vetenskaps-akad. forhandl. Stockholm, 12, pp. 381-388.
'58. Kongliga svenska fregatten Eugenies resa omkring jorden, Zool.,

1, pp. 1-52, 8 pis. Stockholm.
*65. Annulata nova. Ofversigt kongl. yetenskaps-akad. forhandl. Stock-
holm, pp. 22, 167-179, 239-258.
Lord, J. K.

*66. The naturalist in Vancouver Island and British Columbia. 2 vols.,
8vo. London.
M»Int06h, W. C.

'78. Beitrftge zur anatomic von Magelona. Zeitsch. fttrwissensch. zooL,

bd. 31, pp. 401-472, 10 taf.
'85. Report on the Annelida. Polychaeta. Voyage of H. M. S. " Chal-
lenger,'" Zoology, vol. 12, pp. 86+564, 94 pis., and map.
Malaquin, A.

*93. Recherches sur les Syllidiens. Morphologic, anatomic, reproduction,
d^veloppement. 477 pp., 14 pis. Lille: L. Danel.
Malmgren, A. J.

'65. Nordiska Hafs-Annulater. Ofversigt kongl. vetenskaps-akad. for-
handl. Stockholm, 22, pp. 61-110, 181-192, 365-410, 8 tafl.
v. Marenzeller, E^

'79. Stidjapanische Anneliden, 1. Denkschr. k. akad. wissensch. Wien,

math.-naturwissensch. classe, bd. 41, pp. 109-164, 6 taf.
'84. SOdjapanische Anneliden, 2. Denkschr. k. akad. wissensch. Wien,

math.-naturwissensch. classe, bd. 49, pp. 197-224.
'87. *Polychftten der Angra Pequena Bucht. Zool. jahrb., abth. fOr system-

atik. bd. 3, pp. 1-24, 1 taf.
*90. Annulaten des Berings-Meeres. Ann des. k. k. naturhist. Hofmu-
seums, t, 5, pp. 1-8, 1 taf. AV^ien.
Mesnil, F., & M. Caullery.

'd8. Etudes de morphologie exteme chez le.s Ann61ides. 4. La famille
nouvelle des Levinsfenieus, etc. Bull, scient. de la France et de la Bel-
gique, t. 31, pp. 126-160, 1 pi.
Michaelsen, W,

'92. Polychaeten von Ceylon. Jahrb. der Hamburg wissensch. anstalten,
bd. 9, pp. 91-113, Itaf.
Mttller. Fr.

'58. Eiiiiges tiber die Annelidenfauna der Insel Santa Catharina an der
brasilianischen kUste. Arch. fQr naturgesch., jahrg. 24 J1868), pp. 211-
220, 1 taf.
Murdoch, J.

'84. Description of seven new species of Crustacea and one new worm
[Arenicola glacialis'} from arctic Alaska. Proc. U. S. nat. mus., vol.
7, pp. 618-622.

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JOHNSON : POLYCHAETA OF PUGET SOUND BEGION. 437

Roule, L.

*96. Rfisultats scientiflques de la Campagne du "Caudan" — 1896. An-
n^lides, O^phyriens. Ann. de Tuniv. de Lyon, pp. 489-474, 7 pis.
Paris: Masson et Cie.
Stimpson, Wm.

*64. Descriptions of new species of marine Invertebrata from Puget
Sound, collected by naturalists of the Northwest Boundary Commission.
Proc. acad. nat. sciences Phila., 1804, pp. 163-161.
Verrill. A. E.

: 00. Additions to the Turbellaria, Nemertina, and Annelida of the Ber-
mudas, etc. Trans. Conn. acad. arte and sciences, vol. 10, pp. 695-672,
Ipl.
Webster, H. E.

'79. On the Annelida Chaetopoda of the Virginian coast. Trans. Albany
institute, vol. 9, pp. 202-272, 11 pis.
Webster, H. E., & J. E. Benedict.

'87. The Annelida Chaetopoda from Eastport, Maine. Report U. S.
comm. iish and fisheries for 1886-87, pp. 707-766, 8 pis.
Wir6n, A.

*B3. Chaetopoder fr&n Sibiriska Ishafvet och Berings haf , insamlade under
Vega-expeditionen, 1878-79. Vega-expedlt. vetensk. jakttagelser, bd. 2,
pp. 381-428, 6 taf.

Printed, August, 1901.

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Johnson.— Polychmet*.

PLATE 1.'

Fig. 1. Anterior extremity, dirsal aspect, of PAynoe fragilis. The elytra
have fallen ofif. X 8.0.

FigH. 2-7. Harmothoe iphionelloidea.

Fig. 2. Anterior extremity, dorsal aspect ; proboscis exserted ; anterior ely-
tra removed. X 8.6.

Fig. 3. Fifth elytron, right .side. X 8.5.

Fig. 4. Third foot from right side, dorsal aspect. The setae above the dorsal
cirrus all belong to tlie dorsal fascicle, x 28.

Fig. 6. Ventral seta-tip, profile. X 200.

Fig. 0. Stout, curved, dorsal seta. X 200.

Fig. 7. a. Slender doi-sal seta. X •'>3.

b. Tip of same, more magnified. X 200.

^With very few exceptions (in each instance, stated), the tlgares are from camera
fh-awin^fl. In all the plates the drawings have been reduced one-half, and the mA^iflt
cation, as piven with the explanation of each tlKure, has been corrected accordingly. '

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Johnson. — Polyciiaeta.

Plate 1.

M.Pj:^LHj:.del,

Pkoc. Bost. Soc. Nat. Hist. Vol. 29.

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JOHVSOK. " PolychMU.

PLATE 2.

Figs. 8-18. Harmothoe complanata.

Fig. 8. Anterior extremity, dorsal aspect. X 13.

Fig. 9. Twenty-first foot, anterior aspect. X 23.

Fig. 10. Elytron from left side ; the nerves radiating from the elytrophore are

distinctly seen. X 13.

Fig. 11. Tip of stout dorsal seta. X 200.

Fig. 12. Top of slender dorsal seta. X 100.

Fig. 13. Tip of ventral seta, x 200.

Figs. 14-17. Harmotkoe paciflca.

Fig. 14. Anterior aspect of second foot. X 13.

Fig. 16. Tip of slender, slightly curved dorsal seta. X 100.

Fig. 16. Tip of strongly-curved dorsal seta. X 100.

Fig. 17. Tip of ventral seta. X 100.

Fig. 18. Harmothoe tuta ; anterior extremity, after removal of elytra. X 8.5.

Fig. 19. Seventeenth parapod of the same. Anterior aspect. X 17.

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JonNSON. — POLYCHAETA.

Plate 2.

TTM^dU

Proc. Bost. Soc. Nat. Hist. Vol. 29.

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JoHNHON. — Polychaeta.

PLATE 3.

Figs. 20-22. Harmothoe tuta.

Fig. 20. Elytron from left .side. X (».').

Fig. 21. I)on>al seta from eighteintli foot. X 200.

Fig. 22. Ventral seta-tip from eighteenth foot. X 200.

Figs. 2.*)-2.'). Podarke jmgettensis.

Fig. 28. Anterior extremity, probosci.s exserted. X 16.
Fig. 24. Parapod from middle of the body, X 32. o.
Fig. 25. Tip of a ventral (ecmipound) !*eta. X 200.

Figs. 2(^-30. Ntreis virens.

Fig. 20. Anterior extremity, dorsal aspect, of large specimen. X 0.5.

Fig. 27. Foot from middle of body, anterior as] ect. X ^.5.

Figs. 28-30. Tii»s of .**etae from ventral fascicle, sliowing graduation in length

of ap])endage. X 150.
Fig. 31. Tenth foot of Xereis vextllosa, posterior asi ect. X 23.5.
Fig. 32. Natatoiy seta, female Heieronereis of X. vexillosa. X 2tX>.

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Johnson. — Polyciiakta.

Plate 8.

HT.Jy^LH.J^deL

Pboc. Bost. Soc. Nat. Hist. Vol. 29.

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JoHirsoH . — Polycbaeta.

PLATE 4.

Figs. 33-38. Nereis vexillosa.

Fig. 33. Anterior aspect of heteronereized foot. X 8.5.

Fig. 34. Foot from posterior portion of body of young female. X 8.5.

Fig. 35. Foot from p<i8terior region of large adult female, with much elon-
gated dorsal lobe. 6. o. Blood-vessel, ov. Ova lying within coe-
lomic space. X 8.5.

Fig. 30. Stout Hickle-«haped seta from dorsal ramus. X 150.

Figs. 37, 38. Sickle-shaped and " fish-bone" setae from ventral ramus. X 150.

Figs. 3(M5. Nereis agassizi.

Fig. 30. Anterior extremity. X 8.5.

*' Fish-bone " seta from dorsal ramus. X 200.
, 42. Sickle-shaped setae from ventral ramus. X 200.
Hooked seta from doi-sal ramus. X 200.
Foot from middle of series. The glands {g.) at base of upper lobe ju*e

shown. X 23.
Foot from jx^sterior portion of body. X 23.

Anterior, ventral aspect of a young Nereis cyclurus. The prosto-
mium is partially withdrawn into the large, collar-like peristomium.
X 8.5.
Fig. 47. Nereis procern ; foot from anterior portion of body. X 23.5.

Fig.

40.

Figs

.41

Fig.

43.

Fig.

44.

Fig.

45.

Fig.

40.

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JOHN80>f. P0LY( HAKTA.

Plate 4.

HPj:.del.

Puoc. Bust. Soc. Nat. Hist. Vol. 29.

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JomrsoK. " PolycbaeU.

PLATE 6.

Figs. 48-52. NtreU epelurus,

Fig. 48. Anterior extremity, dorsal aepeet, showiag large, coUariform peri-
stomium, within whicli the pro«tomius ia partially withdrawn.
Young specimen. X 8.5.

Fig. 49. Tenth foot, anterior aspect. X 13.

Fig. 50. Falcate seta from lower i^acicle, fentral ramiB, of foot from anterior
region of body. X 200.

Fig. 51. Falcate seta from upper fascicle of ventral ramus. X 200.

Fig. 52. " Fish-bone *' seta, upper fascicle^ rentral ramus. X 200.

Figs. 53-59. Nereis proeera.

Fig. 53. Anterior extremity, doreal ajq>ect. X 8.6.

Fig. 54. Forty -first foot, showing 8tout» dorsal seta and three slender ones.

X23.
Fig. 55. Foot further back ; slender dorsal setae no longer present. X 23.
Fig. 66. Foot from posterior region, near pygidium. X 23.
Figs. 57, 58. '* Fish-bone " and falcate setae from Tentral ramus. X 200.
Fig. 59. 8tout dorsal seta. X 200.

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JOIIXSON. — POLYCHAKTA.

J*LATE 5.

HPJ.del

Proc. Bost. Soc. Nat. Hist. Vol. 29.

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JoHKtOK.— PolxcbAeta.

PLATE 0.

Figs. 60-66. Euphroayne heterobranchia.

Fig. 60. The prostomium, dorsal aspe<rt, showing the antennae, the tentacle, the

dorsal eyes, and the caruncle. X 40.
Fig. 61. Tip of one of the unserrated, cleft, dorsal setae. X 200.
Fig. 62. Tip of serrated, cleft. dor.sal seta. The serrations are almost restricted

to the lower fork. X 200.
a. Tip of another seta of same kind, fully serrated. X 200.
Figs. 63, 64. Bifid, dorsal setae of two forms. X 200.
Fig. 60. Ventral seta of the usual form. X 200.
Fig. *i6 a, 6, c. Various forms of branchiae. X 32.

Figs. 67-70. Pionosyllia elongata.

Fig. 07. Anterior portion of immature specimen (146 somites). X 32.
Fig. 68. Po.sterior extremity, showing pygidiuni,anus(a.),and anal cirri. X 32.
Fi^'. 69. Foot from middle of body. X 62.

Fig. 70. Seta tip. Minute serrations sometimes present on concave side of
falcate appendage. X 275.

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Johnson. — Polychakta.

Plate 6.

HPJ.del

Proc. Bost. Soc. Nat. Hist. Vol. 29.

Digitized by VjOOQ IC

JoHNHON. — rolycbaeta.

PLATE 7.

Fig. 71. PionoayllU dongaia ; outline of anterior portion seen on the ventral
side, showing alimentary canal in situ. The dorsal cirri and the
setae are omitted. p<i. Papillae, t. Tooth, oes. Oesophagus, proo.
Pro ventricul us, or "gizzard.'* c. Coeca. X 28.5.

Figs. 72-76. TrypanoByllU gemmipara.

Fig. 72. Anterior extremity, dorsal aspect, p. Palpus. X 18.

Fig. 73. Proboscis dissected out, showing chitinous teeth and surrounding

fleshy pi^illae (pa.). X 23.5.
Fig. 74. Foot from anterior region of body. The monillform character of the

dorsal cirrus is too strongly indicated, o. c. Ventral cirrus. X 22.5-
Fig. 75. Seta-tip, profile. In glycerine. X 287.
Fig. 76. Caudal buds, seen from ventral aspect The center of proliferation is

on the right, where a cluster of young buds is seen. X 28.

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Johnson. — Polychaeta.

Plate 7.

Proc. Bost. Soc. Nat. Hist. Vol. 29.

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JoHHftosr. — Polyobaeuu

PLATE 8.

Figs. 77-85. Northia elegans.

Fig. 77. Anterior extremity, dorsal aspect. X 6.

Fig. 78. Anterior extremity, specimen lying obliquely on right side. The

minute eyes are visible at base of left articulated antenna, p. c.

Peristomial cirri, p. Palpus, x 8.6.
Pig. 79. Maxillae, somewhat spread apart, and the right edentate maxilla

folded back, to show underlying toothed jaw. x IH.
Fig. 80. Anterior aspect of second foot. br. Gill. d. c. Dorsal cirrus, c. c.

Ventral cirrus. X 13.
Fig. 81. Twenty-eighth foot, profile view. (Lettering as in Fig. 80.) X 13.
Figs. 82, 83. Uncinus or " hooded crotchet," and capillary subulate seta, both

from ventral fascicle of third parapod. X 200.
Fig. 84. Capillary seta with striated border, ventral fascicle of thirty-first foot.

X 200.
Fig. 86. Concave seta (buried in foot) from dorsal fascicle of thirty-first foot.

X 200.
Fig. 86. Anterior extremity, ventral aspect, of Northia iridescens, X 8.5.
Fig. 87. Maxillae of the same, spread widely apart, and seen from dorsal

aspect. X 16.

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Johnson. — Polycii aeta.

Plate' 8.

i H.J? J, del

Proc. Bo8t. Soo. Nat. Hist. Vol. 29.

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J4>UN»«>M. — Folychacta

PLATE 9.

Figs. 88-92. Northia irUUscena.

Fig. 88. Anterior extremity^ dorsal aspect Free-hand drawing, x 3.

Fig. 89. Foot from middle of series, showing the filiform gill and sabolate

dorsal cirrus, x 23.5.
Fig. 90. Wing-tipped uncinos, from a middle parapod. x 200.
Fig. 91. Hooded crotchet from ventral fascicle of third foot. X 190.
Fig. 92. Capillary seta with striated border. X 200.

Figs. 93-100. Lutnbriconereis zonata.

Fig. 93. Anterior extremit>% dorsal aspect X 8.6.

Fig. 94. Maxillae, somewhat spread apart, and seen from the dorsal side.

X 23.6.
Figs. 96, 96. A "spiral" somite from posterior fegion of body, seen from the

dorsal (Fig. 95) and from the ventral aspect (Fig. 96); 1, 2, 3. the

somites involved, x 8.6.
Fig. 97. Fifteenth foot, posterior aspect X 40.
Fig. 98. Foot from middle of series, posterior aspect Unclni only are pres-

ent X 23.6.
Fig. 99. Double-bordered seta from an anterior foot. X 237.
Fig. 100. " Hooded crotchet" from a posterior foot. X 237.

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Johnson. — Polychaeta.

Plate 9.

H.VJ.dcl^

P»o(\ BosT. Soc. Nat. Hist. Vol. 29.

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JoHirsoir. ~ PolychMta.

PLATE 10.

Fig. 101. Posterior aspect of foot of Glycera rugosa, from middle of body
fihowiug 6-lobed gill, and dorsal cirrus {d. c.) seen by transparency
through uppermost lobe of gill (the tip of which has been cut off).
The minute ova have entered the gills, c. c. Ventral cirrus, x 23.i>.

Fig. 102. Jaw and jaw-appendage (ap.) of the same. X 13.

Fig. 108. Posterior aspect of foot of Glycera nana. Dorsal cirrus is not
shown. V. c. Ventral cirrus. X 23.

Fig. 108a. Jaw-appendage of the same. X 40.

Fig. 104. Posterior aspect of a foot from middle of series of He mipodia borealis.
Setae are all compound, and several are almost entirely withdrawn
witliin the foot. The single acicula is indicated by dotted lines.
d. c. Dorsal cirrus, v. c. Ventral cirrus. X 75.

Fig. 104a. Jaw-appendage of the same. The attached end is somewhat ex-
panded. X 75.

Figs. 105-110. Scoloplos elongata.

Anterior extremity, dorsal aspect pp. Palpode. X 13.

Anterior extremity, ventral aspect, with proboscis everted, showing

foliaceous expansions. X 18.
Profile of seventeenth foot, anterior aspect. X 23.6.
Vertical section of foot from middle of length, showing the dorsally-

directed parts, br. Gill. X 52.6.

Portion of one of the deeply-serrated, anterior setae, x 340.
Portion of a ventral seta. X 340.
Anterior extremity of Aricideopsis megalops. X 39.
The same. Foot from posterior portion, x 89.

Fig.

105.

Fig.

106.

Fig.

107.

Fig.

108.

Fig.

100.

Fig.

110.

Fig.

111.

Fig.ai2.

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Johnson. — Polychaeta.

Plate 13.

Proc. Bost. Soc. Nat. Hist. Vol. 29.

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JoHKSox.— Polycbaeta.

PLATE 14.

Figs. 140-142. Ammockares occidentalis.

Fig. 140. ADterior portion, ventral aspect. The tori are represented, but the
uncini are invisible at this magnification. X 13.

Figs. 141 a, b. Uncini, frontal aspect and profile. When in situ, only the por-
tion distal to the constriction is exposed, x 600.

Fig. 142. Seta, showing double serration, x 460.

Fig. 143. Portion of digestive tract of Arenkola claparedei^ showing lower sec-
tion of oesophagus (oes.); oesophageal coeca of two kinds (c, c')« and
the anterior extremity of the chlorogogous tract (cA.). The most
anterior pair of coeca (c.) are always much larger than the rest,
thin-walled and highly vascular. The other coeca (c'.) are thick-
walled and the surface is striated. They are not always paired, and
the number is highly variable (ten in the present instance, which is
about the minimum). Free-hand drawing, x 2.

Fig. 144. Oesophageal coeca of another specimen, showing the thin-walled,
vascular pair (c.) in a much-contracted condition. The smaller coeca
(c'.) number fifteen on each side (with one exception the highest
number hitherto observed) and are <|uite uniform as to size. Several
are attached at or near the median line. Free-hand drawing, x 2.

Figs. 145-148. Cirratulus cingulatus.

Fig. 145. Anterior portion, showing tentacular filament {tf.) and three most

anterior pairs of cirri, all cut short. X 0.
Fig. 14tJ. Two somites from middle of body. The dorsal cirri are broken off,

but the points of attachment are shown. X 13.
Fig. 147. Ventral setae, and two uncini in situ, x 187.
Fig. 148. Ventral hook, middle region, x 150.
Fig. 149. Anterior portion of Cirratulus robustus. The tentacular filament

{t. /.) and dorsal cirri are broken off at point of attachment x 9.
Fig. 150. Somites Si^^S (anterior third of body), left side of the same. Points of

attachment of cirri are distinctly shown, x 13.

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Johnson. — Polychaeta.

Plate 14.

fJUl>\\ f4z.

HM.del.

Paoc. BosT. Soc. Nat. Hist. Vol. 29.

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JoH5tK)N — Polvchaeta

PLATE 16.

Fip*. 161-1 5<?. Pectiiiaria brecicoma.

Fig. 151. Anterior eml, ventral aspect. X 0.

Fig. 162. Seta from ninth somite, with striated border ; slightly twisted. X 100.

Fig. 16.'}. Seta with Htriated border ; not twisted. X 160.

Fig. 164. l^Kiial form of uncinus ; from seventh somite, x 321.

Fig. 166. Occasional form of imcinus ; from twelftJi somite. X 321.

Fig. 166. Scapha-hook, showing laterally-bent tip. X 160.

Figs*. 167-161. SabelUdes anopa.

Fig. 157. Entire /animal, ventro-lateral aspect. Posterior extremity wanting,
as are also most of the tentacles. Free-hand drawing, x 2.6.

Fig. 168. Anterior extremity, right side, tentacles retracted. Two of the
branchiae have been removed from the left side. X 6.

Fig. 169. Single-bordered capillary thoracic seta. X 187.

Fig. 160. Double-bonlered capillary thoracic seta. X 321.

Fig. 161. Uncinus, lateral aHi)ect. X 321.

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ON. — POLYCHAETA.

Plate 15.

Proc. Bost. Soc. Nat. Hist. Vol. 29.

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JoHNBOK.>- Polychaeu.

PLATE 16.

Fig. 102. Anterior extremity of Sabellides anops, dorsal aspect Branchiae on
right side have been cut off at the base. The somites 1-5 are
numbered. X 9.

Fig. 103. Protstomium and tentacleH of the same, dorsal aspect. X 15.

Figs. 104-108. Amphitrite robusta.

Fig. 104. Gill, mounted in glycerine. X 0.

Fig. 106. Seta from thorax. X 187.

Fig. 100. Uncinus from eighteenth somite, in profile. X 321.

Fig. 107. Biserial arrangement of uncinl, thoracic region. X 321.

Fig. 108. Uncinus from near posterior end. X 321.

Figs. 169-171. Amphitrite spiralis.

•

Fig. 109. One of the main branches of a gill, x 15.
Fig. 170. Seta from eighteenth (thoracic) somite. X 321.
Fig. 171 a, b. Uncinl from eighteenth somite, profile. X 321.
Fig. 171 c. Uncinus, anterior aspect, showing five rows of teeth above rostrum.
X 350.

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Johnson. — PoLYniAETA.

Plate 16.

H.PJ.del

Proc. Bost. See. Nat. Hist. Vol. 29.

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JoHxsoM.— Polychaeu

PLATE 17.

Figs. 172-174. Lanice heterobranchia.

Fig. 172. One of the first pair of gills ; ouly one branch ts given in detail. X 9.
Fig. 178. Capillary, double-bordered thoracic seta. X 187.
Fig. 174. Uncinus from nineteenth (thoracic) somite, x 321.

Figs. 175-178. Thelepus crispus.

Fig. 175. Two gill-tilaments, showing place of attachment to body. In glycer-
ine. The bloixl- vessels show through the translucent walls. X 0.
Figs. 17«, 177. Two capillary double-bordered setae of differing form. X loO.
Fig. 178. Uncinus from thirty-second (thoracic) somite, profile view. X 321.
Fig. 178 a, h. Two thoracic uncini of slightly different form. X 350.

Figs. 179-183. Bispira polymorpha.

Fig. 179. Anterior portion, ventral aspect, including thorax and first six so-
mites of abdomen. Free-hand drawing, x 2 (circa).

Fig. 180. Winged seta from dorsal portion of sixth fascicle. X 150.

Fig. 181. Spatulo-mucronate seta-tip from ventral portion- of sixth fascicle. X
276.

Fig. 182. Avicular and mucronate uncini from sixth torus, in their normal re-
lation to each other, x 150.

Fig. 183. Avicular uncinus from abdominal region (twentieth torus), x 150.

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Johnson. — Polychaeta.

Platk 17.

H7fd.del

Proc. Bost. Soc. Nat. Hist. Vol. 29.

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JoHNDOx.— PoIycbaetA.

PLATE 18.

Fig. 184. Portion of gill of Bispira polj/morpha, near base, showing gill-fila-

uientfi and two eyes, x 39.
Fig. 185. Double-bordered, capillary seta from abdominal region. X 150.

Figs. 186-192. Megachone aurantiaccu

Fig. 180. Thorax, and first somite of abdomen, right side. Collar is clearly

indicated, x 0.
Fig. 187. Anterior extremity in end-view, showing regenerating (?) branchiae,

appressed upon the disc ; their tips point ventrad. X 9.
Fig. 188. Gill showing filaments, the blood-vessels of which contain oval blood-

clote. X 26.
Fig. 189. Bordered seta, from thorax. X 187.
Fig. 190. Uncinus from sixth torus, x 150.
Fig. 191. Uncinus from nineteenth torus. X 275.
Fig. 192. Uncinus near posterior extremity. X 275.

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Johnson. — Polychaeta.

Plate 18.

HPd.del

Proc. Bost. Soc. Nat. Hist. Vol. 29.

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JoHNflox. — PolychaeU.

PLATE 19.

Figs. 193-198. Myxicola pacffica.

Fig. 193. Tip of branchia. The web connecting the branchiae is shown as a
border on the right side, and as ruptured tips at w, w. X 12.6.
Double-bordered, thoracic seta. X 450.
Stout, straight seta from thorax, x 460.
Abdominal seta. X 450.
Thoracic uncinus. X 321.
Abdominal oncinus. X 321.

Figs. 199>203. Serpula columbiana.

Profile view of operculum, x 0.
" Bayonet " seta from first fascicle. X 187.
Thoracia seta with striated border, x 187.
Thoracic uncinus (many are d- or 7-toothed). X 275.
Abdominal uncinus. x 275.

Figs. 204-208. Serpula zygophora.

Fig. 204. Chisel-shaped, abdominal seta. X 321.

Fig. 205. Pn)file view of operculum, x 9.

Fig. 20rt. "Bayonet " seu from first fascicle. X 187.

Fig. 207. Thoracic uncinus («-toothed). X 321.

Fig. 208. Abdominal uncinus (usually 6-toothed). X 321.

Fig.

1<H.

Fig.

lltS.

Fig.

196.

Fig.

197.

Fig.

108.

Fig.

1»9.

Fig.

200.

Fig.

201.

Fig.

202.

Fig.

203.

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Johnson. — Polychaeta,

Plate 19.

HPJdel

Proc. Bost. Soc. Nat. Hist. Vol. 29.

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Boston Society of Natural History.

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